GeolCarp_Vol63_No1_49

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA

, FEBRUARY 2012, 63, 1, 49—70 doi: 10.2478/v10096-012-0004-8

Introduction

The  amount  of  geological  and  geochemical  data  from  the
Oligocene—Miocene successions of the North Alpine Foreland
Basin  (NAFB),  northern  Austria,  has  increased  over  the  last
decades, as a number of wells have been drilled for hydrocar-
bon  exploration  (e.g.  Wagner  1998;  Sachsenhofer  &  Schulz
2006). The Oligocene organic-rich sediments were deposited
during  the  early  stages  of  the  Paratethys  separation  and  are
one of the most important sources of hydrocarbons within the
Paratethys  Realm  (Popov  et  al.  2004).  The  Oligocene  sedi-
ments are represented by four formations: Schöneck, Dynow,
Eggerding and Zupfing (Fig. 1).

Three boreholes penetrated the Oligocene sediments of

northern Austria (Sachsenhofer et al. 2010) and proved to be
rich  in  dinoflagellate  cysts  (hereinafter  referred  to  as  di-
nocysts). However, although macro and micropaleontological
studies  have  been  carried  out  extensively  (Cicha  et  al.  1971;
Harzhauser & Mandic 2002; Scherbacher et al. 2002; and ref-
erences therein), the systematic investigation of the dinocysts
has been almost ignored so far except for the pioniering study
of Hochuli (1978) on the Oligocene-Lower Miocene palyno-
morphs  of  the  NAFB,  mostly  focused  on  miospores.  The
present  study  of  dinocysts  was  therefore  undertaken  in  order
to provide additional data for the refining of the stratigraphic
framework of the Oligocene deposits in the NAFB, Austria.

In Central Europe, especially in the Carpathian Flysch,

there are several studies which dealt with the Paleogene di-

Oligocene dinoflagellate cysts from the North Alpine Foreland

Basin: new data from the Eggerding Formation (Austria)

ALI SOLIMAN

Institut für Erdewissenschaften (Geologie und Paläontologie), Karl-Franzens Universität Graz, Heinrichstrasse 26, A-8010 Graz, Austria;

ali.soliman@uni-graz.at

Department of Geology, Faculty of Science, Tanta University, Tanta-31527, Egypt

(Manuscript received January 25, 2011; accepted in revised form June 9, 2011)

Abstract: In spite of detailed geological and geophysical investigations, information available on palynostratigraphy for
the successions deposited in the Austrian part of the North Alpine Foreland Basin (NAFB) is scanty. For the first time,
relatively diverse and well preserved Oligocene dinocyst assemblages, comprising 53 genera and 138 species, are pre-
sented from the organic-rich sediments of the Eggerding Formation. These assemblages contribute to the biostratigraphy
of the Oligocene deposits within the NAFB. Dinocysts such as Chiropteridium lobospinosum, Membranophoridium
aspinatum, Cordosphaeridium spp., Enneadocysta spp., Deflandrea spp., Spiniferites/ Achomosphaera group,
Hystrichokolpoma spp., Apteodinium spp., Glaphyrocysta/Areoligera complex and Wetzeliella spp. represent the main
palynological elements. The occurrence of Chiropteridium spp., Tuberculodinium vancampoae, Distatodinium biffii and
Wetzeliella gochtii is of particular importance for regional correlations within the Lower Oligocene sediments. A compari-
son with age-controlled assemblages from the North Sea Basin, Carpathian and circum-Mediterranean regions substantiate
the attribution to the Rupelian. Lack or sporadic occurrence of the oceanic taxa (e.g. Nematosphaeropsis and Impagidinium)
and dominance of Glaphyrocysta/Areoligera indicate an inner-neritic marine setting during the deposition of the studied
intervals. Although, it is difficult to reconstruct precisely the climatic conditions based on the recorded dinocysts, warm?
sea surface water is suggested. A variation in salinities is interpreted based on the abundances of Homotryblium spp. The
abundance of Peridiniaceae taxa (e.g. Lejeunecysta, Wetzeliella, and Deflandrea) indicates nutrient-rich surface water.

Key words: Oligocene, Austria, North Alpine Foreland Basin, Eggerding Formation, paleoenvironment, dinocysts.

nocysts  and  some  of  them  are  dedicated  to  the  Oligocene.
Gedl (1995) discussed the age and depositional environment
of  the  Ostrysz  Formation  (Oligocene)  of  the  Polish  Inner
Carpathians based on dinocysts. Gedl (2000a,b) investigated
Oligocene  sediments  in  the  Podhale  Paleogene,  Inner
Carpathians,  Poland  and  detected  identical  dinocyst  assem-
blages as in the Eggerding Formation. He used dinocysts and
palynofacies analysis to establish the biostratigraphy and to
reconstruct its paleoenvironment. Gedl (2004a) investigated
the dinocysts at the Eocene/Oligocene boundary at Leluchów,
Poland. Gedl & Leszczyński (2005) examined the deep-ma-
rine Upper Eocene-Lower Oligocene turbiditic and hemipe-
ligic sediments exposed at Folusz, Polish part of the Western
Carpathians.  They  proposed  a  Rupelian  age  for  the  investi-
gated sediments from Magura Beds. Soták et al. (2007) in an
integrated  micropaleontological  study  of  the  Pucov  section,
Podtatranská  Group  used  foraminifera,  calcareous  nanno-
plankton  and  dinocysts  to  detect  the  Eocene/Oligocene  and
Rupelian/Chattian  boundaries.  Recently,  Barski  &  Boja-
nowski (2010) used dinocysts from the Oligocene flysch de-
posits of the Western Carpathians as a tool to investigate the
occurrence  of  the  carbonate  concretions.  Soták  (2010)  dis-
cussed  the  climatic  changes  across  the  Eocene/Oligocene
boundary  in  the  Central-Carpathian  Paleogene  Basin  using
several biota including dinocysts.

Several studies dealing with the Oligocene dinocyst

stratigraphy of adjacent areas have been published from the
North Sea (e.g. Dybkjaer 2004; Schi

ler 2005; Dybkjaer &

SOLIMAN

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

with nannofossils (nannoblooms) occur in the lower part of
the section. This formation is present only in the subsurface
(Wanger 1998).

Material

Twenty-one core samples representing the studied inter-

vals from the Eggerding Formation and the lowermost part
of the Zupfing Formation, obtained from the Puchkirchen 3
(P3), Eggerding 2 (Egdg2) and Oberschauersberg 1 (Osch1)
wells are used in this study (Fig. 2).

In the Eggerding 2 well, the Eggerding Formation is about

14 m thick. The lowermost 1.45 m is composed of sandstone
with  clasts  up  to  10 cm  of  the  Dynow  Formation,  near  its
base. Pelitic rocks intercalating with sandstone beds overlay
the  basal  sandstone.  Remains  of  land  plants  and  fish  occur
frequently  (Sachsenhofer  et  al.  2010).  Four  samples  have
been palynologically investigated from this well; their posi-
tion is shown in Fig. 3.

In the Oberschauersberg Well, the Eggerding Formation is

about 40 m thick and seven samples have been selected for
this study from the lowermost part. The lower part of the Eg-
gerding Formation contains dark grey laminated shaly marl-
stone with white bands of nannoplankton. Fish remains
occur in many samples. The position of the samples is shown
in Fig. 3.

The Eggerding Formation in the Puchkirchen 3 well is

about 20 m thick and composed mainly of dark grey, partly
laminated, marly shales. Fish scales are frequently observed.
The lowermost 30 cm contain clasts of “Lithothamnium
Limestone”, up to 7 cm in diameter. Ten samples from this
well were investigated for dinocysts and their position is
shown in Fig. 3.

Methods

20 g of each sample have been prepared following the

standard  procedures  (e.g.  Green  2001).  The  carbonates  and
silicates were dissolved with HCl and HF, respectively. The
residue was treated for 1 minute in an ultrasonic bath to dis-
aggregate  the  AOM  clusters  before  sieving  at  20 µm.  A
slight  oxidation  with  diluted  HNO

was applied to all sam-

ples. The residue is stained with Safranin “O”. Two to four
slides from each sample were prepared using glycerine jelly
as  mounting  medium,  and  sealed  by  nailvarnish.  The  slides
were studied both qualitatively and quantitatively. The quan-
titative  analysis  included  a  counting  of  the  first  300–or
more–dinocysts  (determinable  and  undeterminable)  when-
ever possible. Freshwater algae and acritarchs recorded dur-
ing  this  process  were  also  counted.  Examination  and
light-photomicrographs  were  taken  using  a  Carl-Zeiss
(Axioplan 2)  light  microscope.  For  photographed  taxa
England Finder coordinates are provided.

Mounts for SEM studies were made by air drying water

suspended residues (re-sieved at 30 m) on glass coverslips
that were mounted on an aluminium stub with a thin-doubled
side sticky tape. Stubs were coated with Platinum. Observa-

Fig. 4. Stratigraphic distributions of dinoflagellate cysts and other
palynomorphs, Egdg2 well, NAFB, Austria.

Series Oligocene
Stage Rupelian
Calcareous nannoplankton

NP23

Formation Eggerding

Fm.

Samples

Egdg2-74 Egdg2-77 Egdg2-83 Egdg2-87

Apteodinium spp.

Caligodinium amiculum

Chiropteridium galea

Cordosphaeridium cantharellus

Distatodinium ellipticum

Glaphyrocysta-Areoligera complex

162

Lejeunecysta spp.

Round brown cysts

Selenopemphix nephroides

Thalassiphora spp.

Wetzeliella spp.

Chiropteridium lobospinosum

Cleistosphaeridium spp.

Cyclonephelium spp.

Dapsilidinium spp.

Deflandrea phosphoritica complex

Enneadocysta pectiniformis

Hemiplacophora semilunifera

Heteraulacacysta porosa

Homotryblium spp.

Hystrichokolpoma cinctum

Impagidinium spp.

Lejeunecysta fallax

Leptodinium italicum

Membranophoridium aspinatum

Operculodinium spp.

Pentadinium goniferum

Polysphaeridium zoharyi

Rhombodinium draco

Selenopemphix armata

Spiniferites pseudofurcatus

Spiniferites/Achomosphaera spp.

Wetzeliella articulata

Wetzeliella asymmetrica

Wetzeliella gochtii

Achilleodinium biformoides

Hystrichokolpoma rigaudiae

Lingulodinium machaerophorum

Batiacasphaera spp.

Cordosphaeridium spp.

Cribroperidinium giuseppei

Cribroperidinium tenuitabulatum

Deflandrea spp.

Distatodinium spp.

Enneadocysta deconinckii

Enneadocysta harrisii

Hystrichostrogylon membraniphorum

Lingulodinium spp.

Palaeocystodinium powellii

Reticulatosphaera actinocoronata

Adnatosphaeridium spp.

Heteraulacacysta campanula

Hystrichokolpoma truncata

Leptodinium membranigerum

Indet. dinoflagellate cysts

Total dinocysts counted

63 451 307 356

Other palynolorphs

Tasmanites spp.

Foraminiferal test linings

Cyclopsiella spp.

Pediastrum spp.

150

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GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

tions  and  photographs  were  made  using  DSM  982  Gemini
SEM,  operating  at  a  working  voltage  of  10 kv.  All  slides,
SEM  stubs  and  residues  are  housed  in  the  paleontological
collection of the Institute of Earth Sciences, Graz University,
Austria. The identified dinocyst nomenclature generally fol-
lows  Dinoflaj 2  available  at  “http://dinoflaj.smu.ca/wiki/
Main_Page”   (Fensome et al. 2008), where the taxonomic
references are cited. Images of the most important dinocysts
are illustrated in Figs. 10—17. A list of the recorded species is
given in Appendix 1.

Results

All samples contain fair to well-preserved palynological

assemblages  consisting  of  dinocysts,  acritarchs,  bisaccate
pollen,  other  pollen  grains,  spores,  microforaminiferal  test
linings and freshwater algae. This study concentrates mostly
on dinocysts as they provide more information to assess the
age of the investigated samples. Although bisaccate pollen is
abundant and spores occur sporadically, no attempt has been
made  to  identify  their  taxonomy.  In  addition,  most  samples
are dominated by organic debris which is mostly composed
of amorphous organic matter (AOM) and terrestrial elements
(opaque and translucent phytoclasts). More details about pa-
lynofacies and its implications is published in Sachsenhofer
et al. (2010). The results of the studied wells are reported in
occurrence-charts  (Figs. 4—6)  with  numbers  representing
specimens counted for each taxon/category. Several age-diag-
nostic  dinocyst  species  are  encountered  and  their  biostrati-
graphic significance is discussed accordingly. The results of
each well will be presented in the following sections.

Eggerding 2 (Egdg2) well

Out of four samples analysed, three are productive and

yielded well preserved dinocyst assemblages. The qualitative
and quantitative composition of the recorded assemblages is
presented in Figures 4 and 7. The assemblage shows a rela-
tively high diversity, as 71 taxa were identified. The Glaphy-
rocysta/Areoligera

complex

(including

Glaphyrocysta,

Areoligera and Cyclonephelium taxa) represents the dominant
group (6.0—45.0 %). Cordosphaeridium spp. (1.4—9.0 %);

Series Oligocene
Stage Rupelian
Calcareous nannoplankton

NP23

Formation Eggerding

Fm.

Samples

ch1-5

ch1-6

ch1-7

Chiropteridium lobospinosum

Diphyes colligerum

Adnatosphaeridium multispinosum

Batiacasphaera sphaerica

Hystrichokolpoma cinctum

Cordosphaeridium cantharellus

2 4

11 73 19 87

Achilleodinium biformoides

Membranophoridium aspinatum

2 2

Apteodinium spp.

Homotryblium spp.

Polysphaeridium zoharyi

2 1

Batiacasphaera micropapillata

Hystrichokolpoma rigaudiae

Hystrichokolpoma truncata

Stoveracysta spp.

Hystrichokolpoma sp. cf. H. salacia

Cyclonephelium spp.

8 1

Impagidinium spp.

Batiacasphaera explanata

34 6 16

3 49 8 3

Hystrichokolpoma spp.

71 1

1 13

Glaphyrocysta/Areoligera complex

153 4 3 43 51

Selenopemphix nephroides

1 1

4 2

Round brown cysts

Deflandrea phosphoritica

1 17 17

Fibrocysta axialis

Hemiplacophora semilunifera

Lingulodinium pycnospinosum

1 2

Distatodinium ellipticum

Distatodinium paradoxum

Spiniferites pseudofurcatus

Cribroperidinium giuseppei

Cribroperidinium tenuitabulatum

Apteodinium spiridoides

Lingulodinium machaerophorum

3 2

Wetzeliella articulata

Enneadocysta pectiniformis

Wetzeliella gochtii

Spiniferites/Achomosphaera spp.

Pentadinium laticinctum/taeniagerum

Chiropteridium galea

13 3

Wetzeliella spp.

14 9

Lejeunecysta spp.

Reticulatosphaera actinocoronata

Adnatosphaeridium robustum

Cordosphaeridium inodes

Deflandrea scabrata

Distatodinium craterum

Enneadocysta arcuata

Enneadocysta harrisii

Leptodinium membranigerum

Thalassiphora spp.

Operculodinium spp.

Wetzeliella asymmetrica

Apteodinium maculatum subsp. grande

Cleistosphaeridium spp.

Enneadocysta deconinckii

Leptodinium italicum

Distatodinium scariosum

Charlesdowniea columna

Dapsilidinium spp.

Heteraulacacysta porosa

Palaeocystodinium golzowense

Rhombodinium draco

Rhombodinium pustulosum

Tuberculodinium vancampoae

Hystrichokolpoma sp. A

Indet. dinoflagellate cysts

Series Oligocene
Stage Rupelian
Calcareous nannoplankton

NP23

Formation Eggerding

Fm.

Samples

ch1-

Total dinocyst counting

323 18 30 193 358 34 281

Other palynomorphs

Tasmanites spp.

3 1

Foraminiferal test lining

Cyclopsiella spp.

Pediastrum spp.

Fig. 5. Stratigraphic distributions of dinoflagellate cysts and other
palynomorphs, Osch1 well, NAFB, Austria.

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GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

Spiniferites/Achomosphaera

com-

plex (including all identified and uni-
dentified  taxa  of  both  genera)
(1.0—28.0 %),  Apteodinium  (1.5—
20.8 %),  Wetzeliella  (10.7—13.4%)
and  Deflandrea  (3.0—10.5 %)  are
fairly  represented.  Low  percentages
of

Operculodinium

(0.2—7.5 %),

Cleistosphaeridium

(1.6—3.0 %),

Dapsilidinium (1.3—2.3 %), Hystri-
chokolpoma (0.6—1.0 %), Rhombo-
dinium

draco

(0.0—1.6 %),

Homotryblium

(1.6—13.0 %)

and

Chiropteridium (1.5—6.4 %), have
also been recorded. Many other taxa
occur sporadically (Fig. 7).

Oberschauersberg 1 (Osch1) well

Seven samples were investigated

and all of them yielded dinocysts. 67
taxa were identified and are quantita-
tively  represented  in  Fig. 5.  The  re-
corded  assemblages  are  similar  to
those from the Eggerding 2 well. Some
taxa  are  abundantly  recorded  in  all
samples, including Deflandrea (1.4—
8.8 %),  Wetzeliella  (4.6—10.0 %),
Hystrichokolpoma

(0.5—28.0 %),

Glaphyrocysta/Areoligera  complex
(5.0—47.0 %)  and  Cordosphaeri-
dium (0.6—31.0 %). Some other taxa
are persistently encountered, for ex-
ample,  Apteodinium  (0.6—3.0 %),
Homotryblium  (0.7—1.4 %),  Penta-
dinium  (0.0—4.6 %),  Polysphaeri-
dium

(0.3—1.0 %),

Spiniferites/

Achomosphaera complex (1.4—3.0 %)
and Stoveracysta (0.7—3.0 %) (Fig. 8).

Puchkirchen 3 (P3) well

84 taxa were identified from this

well,  reflecting  the  highest  diversity
of dinocysts among the studied wells
(Fig. 6).  Apteodinium  (0—17.0 %),
Cordosphaeridium  (1—10.0 %),  De-
flandrea (0.3—35.1 %), Glaphyrocysta/
Areoligera  complex  (1.0—26.0 %),
Spiniferites/Achomosphaera

(1.3—

31.0 %)

and

Homotryblium

(1.0—20.0 %)  and  Wetzeliella  (0.0—
30.0 %)  are  the  most  abundant  taxa.
Figure 9  shows  many  more  taxa  but
with  low  percentages,  for  example,
Cleistosphaeridium,  Hystrichokolpo-
ma,  Impagidinium,  Lingulodinium,
Operculodinium,  Polysphaeridium,
and Thalassiphora.

Fig. 6. Stratigraphic distributions of dinoflagellate cysts and orther palynomorphs, P3 well,
NAFB, Austria. Continued on the next page.

Epoch Oligocene

Stage Rupelian

Chattian

Calcareous nannoplanktons

NP23

NP24

Formation

Eggerding Fm.

Zupfing Fm.

Samples

P3-2

P3-4

P3-6

P3-

Rhombodinium draco

72 3 1

1 2

Deflandrea phosphoritica

63 104 27 106 1 11 36 25 6 2

Homotryblium spp.

9 5 3 1 60 24 5 7

Spiniferites/Achomosphaera spp.

28 9 30 3 4 6 19 38 62 38

Glaphyrocysta/Areoligera complex

15 57 3 35 70 11 13 96 9 11

Cordosphaeridium cantharellus

14 12 14 15 3 4 28 15 4 11

Operculodinium spp.

12 1 5

3 3

Cleistosphaeridium spp.

8 1 4 2 1 6 9 10 14 9

Chiropteridium galea

3 12 6 1

Wetzeliella articulata

6 11 6 11 1 4 9 5

Cordosphaeridium minimum

Wetzeliella asymmetrica

5 6 1 9 2 6 2 3

Hystrichokolpoma truncata

Wetzeliella spp.

6 29 27 65 17 3 30 11 13

Wetzeliella gochtii

3 4 3 5 3 5 4 4

Lingulodinium machaerophorum

3 3 16

Thalassiphora spp.

2 7 3

Lejeunecysta spp.

2 12 73 5 108 26 1 5 5 5

Membranophoridium aspinatum

2 11 1 7 2 1 13 11

Distatodinium craterum

2 1

Adnatosphaeridium multispinosum

23 7

Apteodinium spp.

28 5 4 1

Adnatosphaeridium robustum

Cordosphaeridium fibrospinosum

Lejeunecysta fallax

1 8 5 3 18 6 2 2 1

Distatodinium ellipticum

1 6 1

4 2 23

Polysphaeridium zoharyi

1 3 3 1

4 1 5 12 5

Round brown cysts

1 1 4

24 1 3

6 14

Distatodinium paradoxum

1 1 3 1 1

Batiacasphaera explanata

26 1 3

Lejeunecysta communis

1 2 6 3 6

2 3

Enneadocysta pectiniformis

1 1

Leptodinium italicum

3 5

Melitasphaeridium spp.

Hystrichokolpoma spp.

Heteraulacacysta porosa

1 5

Spinidinium spp.

Operculodinium microtriainum

1 2

Pentadinium goniferum

Deflandrea leptodermata

Dracodinium sp.

Wetzeliella cf. W. ovalis

Lejeunecysta diversiforma

1 6

11 1

Araneosphaera stephanophorum

2 6 2 4

Impletosphaeridium insolitum

Pentadinium laticinctum/taeniagerum

3 8 1

Operculodinium centrocarpum

1 1

Selenopemphix armata

Caligodinium pychnum

Selenopemphix brevispinosa

Reticulatosphaera actinocoronata

3 12

Selenopemphix nephroides

8 1 5 2

2 13

Chiropteridium lobospinosum

Heteraulacacysta campanula

Cyclonephelium paucispinum

Deflandrea scabrata

Achilleodinium biformoides

1 2

Impagidinium spp.

Deflandrea heterophlycta

1 1

Caligodinium amiculum

Batiacasphaera sphaerica

Cordosphaeridium gracile

Lejeunecysta hyalina

1 1

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GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

Age assignments

Age based on dinocysts

However, earlier studies indicated that the Eggerding For-

mation and the lower part of the Zupfing Formation were de-
posited  during  NP23—NP24  (e.g.  Sachsenhofer  et  al.  2010)
the recorded dinocyst assemblages represent additional data
to  the  biostratigraphy  of  the  NAFB  (Figs. 4—6).  In  general,
the samples examined are dominated by dinocysts and other
palynomorphs  (Figs. 10—17). Qualitatively, there is no con-
siderable change in the dinocysts throughout the investigated
samples from the three wells. In order to comment on the age
of the studied samples based on dinocysts, a correlation with
chronostratigraphically calibrated dinocyst events in the ad-
jacent areas will be discussed herein.

Together with a persistent population of long-ranging cos-

mopolitan  representatives  of  Glaphyrocysta,  Areoligera,
Spiniferites,  Operculodinium,  Hystrichokolpoma  and  Cleis-
tosphaeridium some other taxa are particularly useful for age
assignment.  The  age  of  some  marker  taxa  are  discussed  in
the following paragraphs taking into consideration their first
and last occurrences in relation to the nannoplankton stratig-
raphy. Most of these marker taxa are recorded from the three
wells, thus the suggested age could be applied for the studied
intervals.

Epoch Oligocene

Stage Rupelian

Chattian

Calcareous nannoplanktons

NP23

NP24

Formation

Eggerding Fm.

Zupfing Fm.

Samples

P3-2

P3-4

P3-6

P3-

Hemiplacophora semilunifera

Hystrichokolpoma cinctum

1 3 1

Hystrichokolpoma rigaudiae

1 5 2 1

Apteodinium emslandense

Gerlachindinium aechmoorum

Wetzeliella spinosa

Spiniferites pseudofurcatus

5 2

Dapsilidinium spp.

4 1 6 2

Apteodinium spiridoides

Heterosphaeridium heteracanthum

1 1

Cribroperidinium giuseppei

Diphyes colligerum

Deflandrea pachyceros

Pentadinium lophophorum

Palaeocystodinium powellii

1 4

Distatodinium biffii

1 1

Hystrichostrogylon membraniphorum

Nematosphaeropsis labyrinthus

Hystrichokolpoma pusillum

Lejeunecysta sp. 3 of Biffi & Grignani, 1983

Rottnestia borussica

Indet. dinoflagellate cysts

29 8 17 10 8 6 14 22 14 22

Total dinocysts counted

354 300 302 302 300 158 306 367 215 296

Other palynomorphs

Palaeostomocystis sp.

Fungal spore

1 1 2

Tasmanites spp.

2 75

47 17 5 11

Foraminiferal test linning

1 3

Cyclopsiella spp.

Pterospermella spp.

Fig. 6. Continued.

Wetzeliella gochtii is one of the

stratigraphic  markers  for  the  Oli-
gocene  and  its  first  occurrence  is
commonly  used  to  recognize  the
Lower Oligocene (Eldrett et al. 2004
and  references  therein).  Brinkhuis
(1992) detected its lowest occurrence
in  northern  Italy  within  the  Reticu-
latosphaera  actinocoronata  Interval
Zone calibrated with the middle part
of  NP21  Zone  (Lower  Rupelian).
Powell (1992) indicated that the first
occurrence  datum  of  W.  gochtii
marks  the  base  of  dinocyst  biozone
Wgo  (Lower  Rupelian),  which  cor-
responds  to  the  base  of  calcareous
nannofossil  Biozone  NP22.  The
same  event  was  documented  from
the  southern  North  Sea  Basin  (van
Simaeys et al. 2005). The lowest oc-
currence  of  W.  gochtii  delineates  the
Eocene/Oligocene  boundary  in  Le-
luchów, Carpathians Mountains, Po-
land (Gedl 2004a). Torricelli & Biffi
(2001) stated that the W. gochtii oc-
curred  in  the  Lower  Oligocene  Nu-
midian  Flysch  in  Oued  El  Guastal
and  El  Gassaa  sections  in  Tunisia.
Globally, the range of W. gochtii has
been documented from the Early Ru-
pelian to mid-Chattian – lower part
of  subzone  C  of  Zone  D15  (Grad-
stein  et  al.  2004).  In  summary,  ac-

cording to Pross et al. (2010) W. gochtii has a stratigraphic
range from 33.1 Ma to 26.4 Ma (NP22—NP25; Rupelian—
Lower Chattian).

Stoveracysta is encountered only in the Osch1 well (S.

conerae, S. ornata, Stoveracysta sp. 1 of Brinkhuis & Biffi
1993, Stoveracysta sp. 2 of Brinkhuis & Biffi 1993). Pross et
al. (2010) recorded the two informally described Stoveracysta
species of Brinkhuis & Biffi (1993) from the Contessa Barbetti
Road section and correlate their last occurrences to Chron
C12n, i.e. 30.8 Ma (NP23; mid-Rupelian).

Distatodinium biffii is found in two samples (P3-20 and

P3-30) from the Zupfing Formation. This could imply a ?Late
Rupelian-Early  Chattian  age  of  the  Zupfing  Formation  (e.g.
van  Simaeys  et  al.  2005).  Avoiding  the  taxonomic  problem,
due  to  the  few  recorded  specimens,  Pross  et  al.  (2010)  cali-
brate  the  first  occurrences  of  D.  biffii  (s.l.  and  s.s.)  from
Umbria-Marche section in Central Italy to NP24.

Only two specimens of Tuberculodinium vancampoae have

been  recorded  from  the  uppermost  studied  part  of  the  Osch1
well (sample 72). According to Brinkhuis & Biffi (1993) the
first appearance of T. vancampoae can be used as a confirma-
tory  event  for  the  detection  of  the  Reticulatosphaera
actinocoronata Interval Zone of Early Oligocene age at Monte
Cagnero  (central  Italy).  The  same  event  has  been  used  by
Torricelli & Biffi (2001) to recognize the Rupelian age of the
Tabarka section of the Numidian Succession, Tunisia.

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GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

E. pectiniformis defines the top of the dinocyst Zone D14na of
Rupelian age contemporaneously with the first occurrences of
Saturnodinium pansum, not recorded in the current study.
Pross et al. (2010), in their integrated study of the dinocyst
events for the Oligocene in the Western Tethys, recorded dis-
crepancies for the last occurrences of E. pectiniformis, but all
are within the Upper Rupelian—Lower Chattian (NP24).

The first occurrence of Chiropteridium lobospinosum has

been recorded from many sites in northern and western Ger-
many corresponding to Lower Rupelian, dinocyst D13na of
Köthe & Piesker (2007). According to Pross et al. (2010) the
first common occurrence of Chiropteridium spp. (C. lobospi-
nosum) has Rupelian age (NP24).

The co-occurrences of Wetzeliella symmetrica, Wetzeliella

articulata and Areoligera semicirculata in such assemblages
indicate a Rupelian age (see Wilpshaar et al. 1996; Torricelli
& Biffi 2001; Williams et al. 2004; Köthe & Piesker 2007;
Pross et al. 2010).

On the basis of the co-occurrence of the discussed taxa, a

Rupelian age is suggested for the studied intervals of the Eg-
gerding Formation from the three wells. This is consistent
with the age suggested by Sachsenhofer et al. (2010) based
on calcareous nannoplankton (NP23) and partly dinocysts.

Regional and global comparison

A comparison with the published data suggests that di-

nocysts are potentially meaningful for recognizing and corre-
lating the Oligocene of the NAFB, northern Austria with other
regions. Several biostratigraphical studies based on dinocysts
have been carried out Oligocene sediments in different regions
of  the  Mediterranean,  Western  Europe  and  Carpathians  (e.g.
Biffi & Manum 1988; Gedl 1995, 2000a,b; van Simaeys et al.
2004,  2005).  Some  dinocyst  biozonations  of  the  Oligocene
have  been  proposed  by  Brinkhuis  &  Biffi  (1993),  Brinkhuis
(1994), Gedl (2000a,b), Bati & Sancay (2007), and Pross et al.
(2010), such schemes are considered in the current study.

Gedl (2000a,b) proposed four Interval Biozones for the

Oligocene of the Podhale, Inner Carpathians, Poland, name-
ly C. lobospinosum, Wetzeliella sp. A, D. biffii and Glaphy-
rocysta  sp.  A.  Based  on  the  recorded  dinocysts  from  the
NAFB  and  Podhale,  the  investigated  interval  is  identical  to
C. lobospinosum,  Wetzeliella sp. A, biozones. The C.  lobo-
spinosum Interval Zone is defined from the first occurrence
(FO)  of  C.  lobospinosum  to  the  FO  of  Wetzeliella  sp.  A.
Many associated taxa of this zone have been recorded in the
current  study,  including  H.  cinctum,  W.  symmetrica,  W.
gochtii, R. draco, D. phosphoritica, Caligodinium amiculum,
R. actinocoronata and many other taxa. The Wetzeliella sp. A
Interval  Zone  is  defined  from  its  FO  to  the  FO  of  D.  biffii.
Wetzeliella sp. A is not defined clearly in the current study,
so  it  may  be  included  in  Wetzeliella  spp.  But  this  zone  is
characterized by the first occurrence of  Caligodinium sp. A
which  is  identified  herein  as  Gen.  et  sp.  indet.  (Fig. 13.5—7;
Fig. 16.10,11). This species is only recorded from the Osch1
well  which  could  mean  that  the  Osch1  samples  are  younger
than those from the other wells.

In general, the recorded assemblage from the three wells is

equivalent to the Reticulatosphaera actinocoronata (Rac)

Interval Zone, Lower Oligocene of Brinkhuis & Biffi (1993)
from central Italy. This zone is defined from the last occur-
rence  of  Areosphaeridium  diktyoplokum,  which  is  not  re-
corded  in  the  current  samples,  to  the  last  occurrence  of
Glaphyrocysta semitecta. Many taxa, namely, Distatodinium
tenerum, W.  gochtii, T. vancampoae  and  C. amiculum  have
their FOs in this zone and are recorded in the NAFB assem-
blage.

Bati & Sancay (2007) define the Wetzeliella gochtii Inter-

val  Zone  (P-Rp2;  Rupelian,  uppermost  NP23  to  lowermost
NP24)  from  Ebulbahar  and  Keleresdere  sections,  Eastern
Anatolia,  Turkey.  They  used  the  lower  occurrence  of  W.
gochtii to define their zone which include many other taxa,
also recorded in the current study, for example,  T. vancam-
poae, D. biffii, M. aspinatum and D. phosphoritica.

Pross et al. (2010) in their refinement of the magnetostrati-

graphic  calibration  of  dinocyst  events  for  the  Oligocene  of
Western  Tethys  (several  sections  in  central  Italy)  discussed
the first and last occurrences of many Oligocene taxa. Many
of  their  taxa  have  been  recorded  in  the  studied  samples  in-
cluding  W.  gochtii,  G.  semitecta,  H.  pusillum,  Stoveracysta
spp., D. biffii, E. pectiniformis, W. symmetrica and C. lobos-
pinosum. A combination of the first and last occurrences of
these taxa ranging from NP22 to NP24 supports the suggest-
ed age of the studied interval from the NAFB, Austria.

Depositional environment

Over the last decades, many studies indicate that the distri-

bution of dinocysts in Recent sediments is controlled by the
interplay between position relative to shore, water tempera-
ture,  salinity,  depth,  light,  nutrients  and  productivity  (e.g.
Wall et al. 1977; Dale 1996; Marret & Zonneveld 2003). The
(paleo)ecological  preferences  of  some  Oligocene  taxa,
grouped  according  to  their  morphological  similarities,  are
discussed  in  detail  in  earlier  studies  (e.g.  Brinkhuis  1994;
Gedl 2005; Sluijs et al. 2005).

The Eggerding Formation is characterized by dark grey

hemipelagites  and  distal  turbidities.  Such  sediments  are  de-
posited  in  a  near-shore  marine  environment,  which  could
represent favourable marine conditions for many cyst-form-
ing dinoflagellates. Many of the recorded taxa in the investi-
gated  samples  have  been  found  in  a  similar  setting  in  the
Upper  Eocene—Lower  Oligocene  hemipelagites  at  Folusz,
Polish Carpathians (Gedl 2005, for more details).

Although, there is no considerable qualitative change in

the  dinocyst  assemblages  throughout  the  three  wells,  there
are  notable  quantitative  changes  for  some  taxa.  In  the  fol-
lowing paragraphs the prevailing paleoenvironmental condi-
tions  during  the  deposition  of  the  NAFB  Oligocene
sediments in each well are discussed based on the co-occur-
rence of the dinocysts.

Egdg2 well

The absolute dinocyst abundance of three samples from

the Egdg2 well is illustrated in Fig. 7. A near-shore paleoen-
vironment during the deposition of this interval could be pre-

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GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

dicted from the abundance of Glaphyrocysta/Areoligera spp.
(Brinkhuis  1994).  The  Peridiniaceae  taxa  as  Wetzeliella  and
Deflandrea show a considerable abundance. Their occurrence
refers  to  rich-nutrient  environments  (Brinkhuis  et  al.  1992).
Spiniferites/Achomosphaera spp. are regarded as representing
a  wide  range  of  depositional  settings  (e.g.  Dale  1996).  Their
high percentages are recorded when the heterotrophic taxa (es-
pecially,  Wetzeliella)  are  at  a  minimum  (sample  Egdg2-83).
This indicates oligotrophic surface water (Vink et al. 2000).

In sample Egdg2-77, Homotryblium shows a considerable

abundance  but  it  is  depleted  in  the  other  samples  (Fig. 7).
Homotryblium  is  broadly  considered  to  be  characteristic  of
restricted  conditions  with  abnormal  salinity  (Brinkhuis
1994).  Pross  &  Schmiedl  (2002)  proposed  temporary  high-
salinity  conditions  during  the  deposition  of  the  Lower  Oli-
gocene  sediments  of  the  Mainz  Embayment  based  on  a
dinocyst  assemblage  dominated  by  Homotryblium  tenuispi-
nosum. The same conclusion was suggested by Köthe (1990,
2009) where intervals of high Homotryblium abundances in
the  Oligocene  of  northwest  and  central  Germany  indicate
high-salinity conditions. It is worthy noting that the acme of
Homotryblium is associated with abundance of the brackish-
water algae Pediastrum (Fig. 4) which implies that Homotry-
blium  forming-cyst  can  tolerate  a  wide  range  of  salinities
(Sluijs  et  al.  2005).  Thus,  salinities  are  verified  during  the
deposition  of  the  lower  part  of  the  Eggerding  Formation
based on the variation of Homotryblium and

O (Sachsen-

hofer  et  al.  2010).  In  addition,  the  occurrence  of  Thalassi-
phora  pelagica  is  distinguished  when  Homotryblium  is
depleted. Pross & Schmiedl (2002) interpreted alternation of
Homotryblium  and  T.  pelagica,  as  an  indication  of  alterna-
tions between high- and low-salinity conditions, respective-
ly.  Williams  &  Bujak  (1977)  have  suggested  that  elevated
abundances  of  Homotryblium  spp.  may  be  associated  with
warm  climatic  conditions.  In  addition,  Sachsenhofer  et  al.
(2010)  concluded  on  the  basis  of  palynofacies  analysis  that
the  Eggerding  Formation  was  deposited  in  an  anoxic  envi-
ronment. The occurrence of Thalassiphora, a genus found in
oxygen-depleted  environments  (Pross  2001),  confirms  this
hypothesis.

Osch1 well

Due to low recovery of dinocysts in the three samples of

the Osch1 well, only the absolute abundance of four samples
is presented in Figure 8. The low abundance of Spiniferites/
Achomosphaera in this well is notable while the high abun-
dance of Hystrichokolpoma spp. is recognizable. Hystricho-
kolpoma may occur in a wide range of marine environments
and has a global distribution (Brinkhuis 1994). Van Mourik
et  al.  (2001)  and  Rasmussen  et  al.  (2003)  considered
Hystrichokolpoma as an open marine indicator. There is also
an inverse correlation between Cordosphaeridium, mostly C.
cantharellus,  and  Glaphyrocysta/Areoligera  spp.  The  latter
is  accepted  as  representing  a  shallow-water  environment
while Cordosphaeridium is associated with open marine wa-
ter masses (Brinkhuis 1994; Rasmussen et al. 2003). In sam-
ple  Osch1-65,  there  is  an  acme  of  heterotrophic  taxa
(Lejeunecysta,  Deflandrea  and  Wetzeliella)  which  indicate

elevated nutrient-rich environment (Pross & Schmiedl
2002).

It is widely accepted that the Early Oligocene is a period of

cold climatic conditions. This is mostly reflected by the abun-
dance  of  the  cold  water  dinocyst  Svalbardella  cooksoniae.
Śliwińska & Heilmann-Clausen (2011) revealed that S. cook-
soniae is present in the narrow interval of Chron 12r, close to
the  NP21/NP22  boundary  in  many  high  and  mid  latitude
Northern  Hemisphere  sections,  ranging  from  the  Greenland
Sea in the North to Italy in the South. S. cooksoniae is not re-
corded in the studied samples which mean that the studied in-
terval  may  be  younger  than  this  event.  On  the  other  hand,
Glaphyrocysta/Areoligera  spp.  dominated  all  studied  sam-
ples. According to Köthe (1990) its abundance indicates warm
climatic conditions. The elevated abundances of Homotryblium
spp. as in Egdg2-77 and P3-16 may be associated with warm
conditions  (Brinkhuis  1994).  Diphyes  colligerum  is  sporadi-
cally  recorded  from  the  studied  samples,  its  occurrence  sug-
gests  warm-water  conditions  (van  Mourik  et  al.  2001;  Gedl
2004b). However, the warm climatic conditions still doubtful
based on the encountered dinocyst assemblage.

It is worth to mention that the occurrences of Impagidinium

(0.7 to 3.5 %) and outer neritic to oceanic indicators, and
Stoveracysta (e.g. Wall et al. 1977; Clowes 1985; Brinkhuis
& Biffi 1993) are approximately contemporaneous (Fig. 8),
which could indicate a short invasion of open marine water.

P3 well

The absolute abundance of dinocysts in P3 well is presented

in Fig. 9. At a glance, Deflandrea  spp. and other (proto)peri-
dinioid  (heterotrophic)  as  Wetzelielloideae  (e.g.  Wetzeliella)
and  Congruentidioideae  (e.g.  Lejeunecysta)  are  abundant  in
the  lower  and  middle  part  of  the  investigated  interval.  Their
co-occurrence indicates nutrient-rich near-shore environments
(e.g.  Pross  &  Schmiedl  2002)  but  some  studies  attribute  the
abundance  of  Deflandrea  to  nutrient  availability  rather  than
distance to the shoreline (e.g. Brinkhuis 1994; Gedl 2005). In
the upper part, the autotrophic taxa such as Spiniferites/Acho-
mosphaera, Homotryblium and Distatodinium show relatively
high abundances. Thus the abundance of the autotrophic taxa
in samples P3-15 to P3-30 could indicate depletion in the nu-
trient supply. Additionally, the co-occurrence of Homotryblium
spp. and Polysphaeridium zoharyi, a near-shore and high sa-
linity  indicator,  indicates  a  change,  probably,  to  a  more  sa-
line  environment  (Brinkhuis  1994;  Gedl  1995;  Dybkjaer
2004; Sluijs et al. 2005). The occurrence of the inner to outer
neritic  genus  Adnatosphaeridium  spp.  matches  Spiniferites/
Achomosphaera  in  the  upper  part  of  this  well  (Fig. 9)
(Brinkhuis & Biffi 1993).

Additional evidence of the near-shore environments are

the  abundance  of  Glaphyrocysta-Areoligera  complex  (e.g.
Areoligera, Cyclonephelium, Glaphyrocysta) (e.g. Brinkhuis
&  Zachariasse  1988)  and  the  considerable  occurrence  of
Membranophoridium  aspinatum  (Gedl  2005).  Normal  ma-
rine shallow water could be indicated by the presence of Im-
pletosphaeridium in the lower part of the well (Islam 1984)
(Fig. 9).  This  conclusion  is  also  supported  by  the  rare  occur-
rence of the open marine genus Impagidinium and Leptodinium.

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GEOLOGICA CARPATHICA, 2012, 63, 1, 49—70

Summarily, the studied interval of the P3 well was depos-

ited, in general, under a shallow-water environment which is
more nutrient-rich in the lower part (samples 2 to 16) than
the upper part.

Conclusions

The following conclusions are drawn from the data dis-

cussed about the Eggerding Formation of the North Alpine
Foreland Basin (NAFB) in Austria:

1. 138 species belonging to 53 genera of dinocysts have

been identified and documented from the Lower Oligocene
of the NAFB, Austria (Appendix 1). Such high diversity of
dinocysts encourages further studies of the Oligocene sedi-
ments in the NAFB which could enhance the biostratigraphic
resolution and paleoenvironmental interpretation as well.

2. Many Oligocene marker dinocysts are recorded, but the

occurrence  of  Chiropteridium  spp.  (C.  lobospinosum),  Fibro-
cysta  axialis,  Tuberculodinium  vancampoae  and  Wetzeliella
gochtii indicate a Rupelian age of the studied interval of the Eg-
gerding  Formation.  The  occurrence  of  Distatodinium  biffii  in
the Zupfing Formation, probably indicates the Lower Chattian.

3. A marine inner-neritic setting could be suggested on the

basis of the encountered dinocysts. The dominance of AOM,
phytoclasts reflect the influence of the terrestrial freshwater
discharge on the area of deposition.

4. A doubtful warm climatic conditions is suggested.
5. The variable occurrences of Homotryblium, P. zoharyi

and pediastrum suggest changes in sea surface salinities dur-
ing the deposition on the studied interval.

6. The co-occurrence (proto)peridinioid (heterotrophic)

taxa indicates nutrient-rich surface water during the deposi-
tion of the Lower Oligocene in the NAFB.

Taxonomic notes

The species presented herein are listed in Appendix (1)

and some of them including marker taxa are illustrated in
Figs. 10—17. Remarks or comments are given for some taxa
and a brief description is provided for dinocysts that are pre-
sented in open or informal nomenclature. Taxa are presented
alphabetically following Fensome et al. (1993).

Division: Dinoflagellata (Bütschli 1885) Fensome et al., 1993

Class: Dinophyceae Pascher, 1914

Subclass: Peridiniphycidae Fensome et al., 1993

Order: Gonyaulacales Taylor, 1980

Genus: Batiacasphaera Drugg, 1970

Batiacasphaera explanata (Bujak in Bujak et al., 1980)

Islam, 1983

Fig. 13.3,4; Fig. 17.2,3

Chytroeisphaeridia explanata Bujak et al., 1980, pl. 13, figs. 13—14
Batiacasphaera explanata (Bujak in Bujak et al. 1980) Islam, 1983, p. 235
Batiacasphaera? sp. 1 Schi

ler, 2005, Pl. 7, fig. 12

R e m a r k s : Bujak in Bujak et al. (1980) described Batia-

casphaera (Chytroeisphaeridia) explanata from the Eocene
of Southern England. In his original description he attributed
a smooth to chagrinate autocyst with spherical to ovoidal
shape and apical archaeopyle to this new species. Recently,
Schi

ler (2005: Pl. 7, fig. 12) attributed a similar morpho-

type from the base of the Oligocene of North Sea but a little
bit longer than broader to Batiacasphaera? sp. 1. It may be
possible that the current material and that of Schi

ler are

conspecific with Bujak’s, if the length/width ratio is not an
important feature (Schi

ler, personal communications).

Thus, all similar morphotypes recorded from the studied
samples are attributed to B. explanata.

Genus: Hystrichokolpoma Klump, 1953 emend. Foucher, 2004

Type species: Hystrichokolpoma cinctum Klumpp, 1953.

Hystrichokolpoma sp. cf. H. salacium Eaton, 1976

Fig. 14.3

Hystrichokolpoma salacia Eaton, 1976, p. 271—272, pl. 11, figs. 1—3,

text-figs.16A—B

R e m a r k s : The recorded specimens are questionably at-

tributed to H. salacium because the SEM investigation
shows processes with thicker striations. Some processes in-
clude small or weakly developed tubuli.

Hystrichokolpoma spp.

Fig. 11.1—4,15,16,18; Fig. 14.6—8

R e m a r k s : A group of different morphotypes attributed to

Hystrichokolpoma based on the presence of two types of pro-
cesses (large and cylindrical in the post and pre-paracingular
area and small in the paracingular and parasulcal areas).

Genus et sp. indet.

Fig. 13.5-7; Fig. 16.10,11

Caligodinium? sp. A, Gedl 2000, p. 231, Fig. 11h, n, o, q, r

R e m a r k s : A holocavate cyst, proximochorate, subspher-

ical to ovoidal in shape, autophragm granular and ornamented
by penitabular rods of equal height supporting an ecto-
phragm across the intratabular area. Apical (rarely seen) and
antapical horns are prolonged from the ectophragm. Sutural
ridges probably delineating a gonyaulacacean tabulation.
The archeopyle is apical and operculum is free.

C o m p a r i s o n : Genus et sp. indet. and Gardodinium Al-

berti 1961; emend. Harding 1996 are similar in having a thin
ectophragm, but the former differs in lacking the apical boss
in the endocyst and the rods supporting ectophragm are pen-
itabular rather than covering the intratabular areas. It differs
from Stoveracysta by having an ectophragm rather than an
autophragm cyst (Clowes 1985). It differs from Alisocysta,
Schematophora and Eisenackia in the intratabular areas cov-
ered by ectophragm. The attribution of similar morphotypes
from the Oligocene sediment of Podhale as Caligodinium sp. A
(Gedl 2000) is rejected due to the holocavate nature.

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Acknowledgment:  Reinhard  Sachsenhofer  (Leoben)  and
Rohöl-Aufsuchung  AG  (RAG)  are  thanked  for  the  well  data
and samples. I am deeply indebted to Drs. W. Piller (Graz) and
S.  Torricelli  (Milan)  for  their  helpful  comments  on  the  early
version. I gratefully acknowledge the constructive reviews of
P. Gedl (Kraków) and the two anonymous reviewers. This work
is supported by the ÖWA and FWF (Project No. 21414-B16).

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Appendix 1

List of the identified dinocyst from the NAFB, Austria (for full taxonomic references see Fensome et al. 2008).

Achilleodinium biformoides (Eisenack, 1954) Eaton, 1976
Achomosphaera alcicornu (Eisenack, 1954) Davey & Williams, 1966
Achomosphaera ramulifera (Deflandre, 1937) Evitt, 1963
Adnatosphaeridium multispinosum Williams & Downie, 1966
Adnatosphaeridium robustum (Morgenroth, 1966) De Coninck, 1975
Apteodinium australiense (Deflandre & Cookson, 1955) Williams, 1978
Apteodinium emslandense (Gerlach 1961) Stover & Evitt, 1978;

emend. Benedek & Sarjeant, 1981

Apteodinium maculatum ssp. grande (Cookson & Hughes, 1964) Below,

1981

Apteodinium spiridoides Benedek, 1972
Apteodinium? vescum Matsuoka, 1983

Araneosphaera stephanophora (Benedek, 1972) emend. Benedek &

Sarjeant, 1981

Areoligera coronata (O. Wetzel, 1933) Lejeune-Carpentier, 1938
Batiacasphaera explanata (Bujak in Bujak et al., 1980) Islam, 1983
Batiacasphaera micropapillata Stover, 1977
Batiacasphaera sphaerica Stover, 1977
Caligodinium amiculum Drugg, 1970
Caligodinium pychnum Biffi & Manum, 1988
Charlesdowniea columna (Michoux, 1988) Lentin & Vozzhennikova,

1990

Chiropteridium galea (Maier, 1959) Sarjeant, 1983
Chiropteridium lobospinosum Gocht, 1960

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Chiropteridium spp.
Cleistosphaeridium ancyreum (Cookson & Eisenack, 1965) Eaton et

al., 2001

Cleistosphaeridium placacanthum (Deflandre & Cookson, 1955) Eaton

et al., 2001

Cordosphaeridium cantharellus (Brosius, 1963) Gocht, 1969
Cordosphaeridium fibrospinosum Davey & Williams, 1966
Cordosphaeridium gracile (Eisenack, 1954) Davey & Williams, 1966
Cordosphaeridium inodes (Klumpp, 1953) Eisenack, 1963 emend.

Sarjeant, 1981

Cordosphaeridium minimum (Morgenroth, 1966) Benedek, 1972

Cribroperidinium giuseppei (Morgenroth, 1966)

Cribroperidinium tenuitabulatum (Gerlach, 1961) Helenes, 1984
Cyclonephelium compactum Deflandre & Cookson, 1955
Cyclonephelium paucimarginatum Cookson & Eisenack ,1962
Cyclonephelium paucispinum Davey, 1969
Cyclonephelium vannophorum Davey, 1969
Dapsilidinium pseudocolligerum (Stover, 1977) Bujak et al., 1980
Dapsilidinium simplex Bujak et al., 1980
Dapsilidinium spp.
Deflandrea heterophlycta Deflandre & Cookson, 1955
Deflandrea leptodermata Cookson & Eisenack, 1965
Deflandrea? pachyceros Deflandre & Cookson, 1955
Deflandrea phosphoritica Eisenack, 1938
Deflandrea phosphoritica var. spinulosa Alberti, 1959
Deflandrea scabrata Wilson, 1988
Deflandrea truncata Stover, 1974
Diphyes colligerum (Deflandre & Cookson, 1955) Cookson, 1965

emend. Goodman & Witmer, 1985

Distatodinium biffii Brinkhuis Powell & Zevenboom, 1992
Distatodinium craterum Eaton, 1976
Distatodinium ellipticum (Cookson, 1965) Eaton, 1976
Distatodinium paradoxum (Brosius, 1963) Eaton, 1976
Distatodinium scariosum Liengjarern et al., 1980
Distatodinium tenerum (Benedek, 1972) Eaton, 1976; emend. Benedek

& Sarjeant, 1981

Dracodinium sp.
Enneadocysta deconinckii Stover & Williams, 1995
Enneadocysta harrisii Stover & Williams, 1995
Enneadocysta arcuata (Eaton, 1971) Stover & Williams, 1995
Enneadocysta pectiniformis (Gerlach, 1961) Stover & Williams, 1995
Fibrocysta axialis (Eisenack, 1965) Stover & Evitt, 1978
Gerlachidium aechmophorum (Benedek, 1972) Benedek & Sarjeant,

1981

Glaphyrocysta espiritosantensis (Regali et al., 1974) Arai in Fauconnier &

Masure, 2004

Glaphyrocysta exuberans (Deflandre & Cookson, 1955) Stover &

Evitt, 1978; emend. Sarjeant, 1986

Glaphyrocysta intricata (Eaton, 1971) Stover & Evitt, 1978
Glaphyrocysta microfenestrata (Bujak, 1976) Stover & Evitt, 1978
Glaphyrocysta ordinata (Williams & Downie, 1966) Stover & Evitt, 1978
Glaphyrocysta retiintexta (Cookson, 1965) Stover & Evitt, 1978
Glaphyrocysta semitecta (Bujak, 1980) Lentin & Williams, 1981
Glaphyrocysta texta (Bujak, 1976) Stover & Evitt, 1978
Glaphyrocysta wilsonii Kirsch, 1991
Glalphyrocysta dentata Matsuoka, 1984
Hemiplacophora semilunifera Cookson & Eisenack, 1965
Heteraulacacysta campanula Drugg & Loeblich Jr., 1967
Heteraulacacysta porosa Bujak in Bujak et al., 1980
Heterosphaeridium heteracanthum (Deflandre & Cookson, 1955)

Eisenack & Kjellström, 1972

Homotryblium aculeatum Williams, 1978
Homotryblium floripes (Deflandrea & Cookson, 1955) Stover, 1975
Homotryblium oceanicum Eaton, 1976
Homotryblium pallidum Davey & Williams, 1966
Homotryblium abbreviatum Eaton, 1976
Homotryblium plectilum Drugg & Loeblich Jr., 1967
Homotryblium tenuispinosum Davey & Williams, 1966

Homotryblium spp.
Homotryblium vallum Stover, 1977
Hystrichokolpoma truncata Biffi & Manum, 1988
Hystrichokolpoma cinctum Klumpp, 1953
Hystrichokolpoma pusillum Biffi & Manum, 1988
Hystrichokolpoma rigaudiae Deflandre & Cookson, 1955
Hystrichokolpoma salacia Eaton, 1976
Hystrichokolpoma sp. A
Hystrichokolpoma torquatum Damassa, 1979
Hystrichostrogylon membraniphorum Agelopoulos, 1964
Impagidinium brevisulcatum Michoux, 1985
Impagidinium dispertitum (Cookson & Eisenack, 1965) Stover & Evitt,

1978

Impagidinium maculatum sensu Schioler, 2005
Impagidinium pallidum Bujak, 1984
Impletosphaeridium insolitum Eaton, 1976
Lejeunecysta communis Biffi & Grignani, 1983
Lejeunecysta diversiforma (Bradford, 1977) Artzner & Dörhöfer, 1978
Lejeunecysta fallax (Morgenroth, 1966) Artzner & Dörhöfer, 1978

emend. Biffi & Grignani, 1983

Lejeunecysta hyalina (Gerlach, 1961) Artzner & Dörhöfer, 1978
Lejeunecysta sp. 3 of Biffi & Grignani, 1983
Lejeunecysta tenella (Morgenroth, 1966) Wilson & Clowes, 1980
Leptodinium italicum Biffi & Manum, 1988
Leptodinium membranigerum Gerlach, 1961
Lingulodinium brevispinosum Matsuoka & Bujak, 1988
Lingulodinium machaerophorum (Deflandre & Cookson, 1955) Wall, 1967
Lingulodinium pycnospinosum (Benedek, 1972) Stover & Evitt, 1978
Melitasphaeridium spp.
Membranophoridium aspinatum Gerlach, 1961
Nematosphaeropsis labyrinthus (Ostenfeld, 1903) Reid, 1974
Operculodinium centrocarpum (Deflandre & Cookson, 1955) Wall,

1967

Operculodinium microtriainum (Klumpp, 1953) Islam, 1983
Operculodinium spp.
Operculodinium tiara (Klumpp, 1953) Stover & Evitt, 1978
Palaeocystodinium golzowense Alberti, 1961
Palaeocystodinium powellii (Strauss et al., 2001)
Pentadinium goniferum Edwards, 1982
Pentadinium laticinctum Gerlach, 1961
Pentadinium taenigerum Gerlach, 1961
Polysphaeridium zoharyi (Rossignol, 1962) Bujak et al., 1980
Pyxidinopsis fairhavenensis de Verteuil & Norris, 1996
Reticulatosphaera actinocoronata (Benedek, 1972) Bujak &

Matsuoka, 1986

Rhombodinium draco Gocht, 1955
Rhombodinium pustulosum Chateauneuf, 1980
Rottnestia borussica (Eisenack, 1954) Cookson & Eisenack, 1961
Selenopemphix armata Bujak, 1980
Selenopemphix brevispinosa Head et al., 1989
Selenopemphix nephroides Benedek, 1972
Spinidinium sp.
Spiniferites crassivariabilis Strauss et al., 2001
Spiniferites pseudofurcatus (Klumpp, 1953) Sarjeant, 1970 emend.

Sarjeant, 1981

Spiniferites ramosus (Ehrenberg, 1838) Mantell, 1854
Stoveracysta conerae Biffi & Manum, 1988
Stoveracysta ornata (Cookson & Eisenack, 1965) Clowes, 1985
Tectatodinium pellitum Wall, 1967
Thalassiphora patula (Williams & Downie, 1966) Stover & Evitt, 1978
Thalassiphora pelagica (Eisenack, 1954) Eisenack & Gocht, 1960
Tuberculodinium vancampoae (Rossignol, 1962) Wall, 1967
Wetzeliella articulata Eisenack, 1938
Wetzeliella ovalis Eisenack, 1954
Wetzeliella echinosuturata Wilson, 1967
Wetzeliella gochtii Costa & Downie, 1976
Wetzeliella spinulosa Wilson, 1988
Wetzeliella symmetrica Weiler, 1956