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STUDIA
GEOLOGICA
POLONICA
Vol. 124, Kraków 2005, pp. 371–394.
Methods and Applications in Micropalaeontology
Edited by J. Tyszka, M. Oliwkiewicz-Miklasiñska,
P. Gedl & M. A. Kaminski
Przemys³aw GEDL1
In situ and recycled dinoflagellate cysts from Middle
Miocene deposits at Bêczyn, Carpathian Foredeep, Poland
(Figs 1–12; App. 1, 2)
Abstract. Middle Miocene clays exposed at Bêczyn (Carpathian Foredeep) contain frequent organicwalled dinoflagellate cysts. The age-assessment of the dinocysts from the clays at Bêczyn suggests
their Early Badenian age (interregional dinocyst zone D17). However, dinocyst assemblage is in large
part composed of recycled pre-Miocene specimens. Late Cretaceous and Palaeogene dinocysts are the
most frequent among the forms treated as recycled. Occurrence of so frequent recycled dinocysts
points at erosion of pre-Miocene substratum during the Early Badenian.
Key words: Miocene, biostratigraphy, dinocysts, Carpathians, Carpathian Foredeep.
INTRODUCTION
The Middle Miocene clays at Bêczyn (former: Benczyn) have been found by
Ksi¹¿kiewicz during his geological mapping of the Wadowice vicinity in the thirties and late forties of the twentieth century (see Ksi¹¿kiewicz, 1951). Since then,
palaeontological investigations carried on the clays at Bêczyn revealed occurrence
of frequent fossils. Krach (in Krach & Ksi¹¿kiewicz, 1948) described over 100 taxa
of Gastropoda and 37 species of Lamellibranchiata (he also found foraminifera,
Bryozoa, Scleractinia, and fish, Arthropoda and Cnidaria remains). Krach’s material served for further palaeontological studies: Ma³ecki (1949) presented a list of
determined Bryozoa taxa, Moenke (1953) gave a detailed description of Scleractinia, whereas £uczkowska (1957) studied foraminifera (she also investigated additional samples collected from the streambed at Bêczyn). Schiller (1976) investigated ostracods from the Bêczyn clays.
1
Institute of Geological Sciences, Polish Academy of Sciences, Senacka 1, 31-002 Kraków,
Poland; E-mail: [email protected]
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Fig. 1.
P. GEDL
Location of Bêczyn (arrowed), Carpathian Foredeep, Poland
Organic-walled dinoflagellate cysts (hereafter dinocysts), being the subject of
present study, have been described so far from more northern parts of the Carpathian Foredeep (Gedl, 1996, 1997, 1999). The purpose of this study was investigation of their assemblages from present-day southernmost part of the Foredeep.
Dinocyst occurrence in this part of the basin may bring new information on palaeogeography of Carpathian Foredeep marine basin during Middle Miocene.
GEOLOGICAL SETTING
The clays that crop out at Bêczyn (Fig. 1) represent Middle Miocene deposits of
the Carpathian Foredeep that have been tectonically disturbed by the Carpathian
overthrust. They plunge at an angle of approximately 40° under the flysch deposits
of the so-called “outer flysch” (Ksi¹¿kiewicz in Krach & Ksi¹¿kiewicz, 1948). This
tectonic unit, recently believed to represent the Skole nappe (see Wójcik et al.,
1999), consists of Palaeogene Ciê¿kowice sandstone and Menilite beds (Ksi¹¿kiewicz in Krach & Ksi¹¿kiewicz, 1948; Ksi¹¿kiewicz, 1951). According to Ksi¹¿kiewicz (in Krach & Ksi¹¿kiewicz, 1948) the clays at Bêczyn rest upon the Jurassic of
the Carpathian foreland, whereas the younger Miocene units cover the folded Carpathian nappes (see also Nowak, 1959; Wójcik et al., 1999).
The hitherto palaeontological age-assessments of the clays at Bêczyn suggest
their Badenian or early Badenian (Moravian) age. The authors of early papers
(Krach in Krach & Ksi¹¿kiewicz, 1948; Ma³ecki, 1949) when describing the clays
at Bêczyn referred to lower Tortonian, i.e., the out-of-date equivalent of the lower
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
Fig. 2.
373
Location of investigated samples of Middle Miocene clays at Bêczyn
Badenian, a stage in chronostratigraphic division of the Central Paratethys (see
Papp et al., 1978).
MATERIAL
The investigated clays are poorly exposed in the streambed of a small creek at
Bêczyn (Fig. 2) – their exposures are situated mainly in creek meanders (Fig. 3A).
These sediments consist of greenish and greyish clays with infrequent layers of reddish clays and sandy clays. The clays lack well-marked stratification. Local massoccurrences of macrofauna detritus, obviously reworked from shallower environments, form layers and lenses, which dip 40–45° southwards (Fig. 3B).
Seventeen samples were collected from the clays exposed at Bêczyn (Fig. 2; Appendix 1). They were taken either from clays with abundant macrofauna as well
from those devoid of shell detritus. Investigated samples were numbered consequently from the northernmost sample (Bnz1) to sample Bnz17 collected in southernmost limits of the Bêczyn clay exposures, close to the Carpathian overthrust
(Fig. 2). Their relation is not certain due to the lack of continuous exposure and uncertain tectonic disturbance. However, because of the general southward dip of
strata (also mentioned by Ksi¹¿kiewicz in Krach & Ksi¹¿kiewicz, 1948), sample
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P. GEDL
Fig. 3. Middle Miocene clays in the streambed at Bêczyn. A – typical exposure in a stream
meander; B – 40–45° dipping southwards shellbed with the position of samples Bnz13 and Bnz14
(coin diameter – 23 mm)
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
375
Bnz1 is tentatively treated as the oldest one, whereas the remaining samples are
treated as arranged in stratigraphic order, with the youngest sample being Bnz17.
METHODS
The samples were processed following a standard palynological procedure including 38% hydrochloric acid (HCl) treatment, 40% hydrofluoric acid (HF) treatment, heavy liquid (ZnCl2+HCl; density 2.0 g/cm3) separation, ultrasound for
10–15 s and sieving at 15 µm on a nylon mesh. The quantity of rock processed was
30 g. Two microscope slides were made from each sample using glycerine jelly as a
mounting medium. Dinocysts were counted from both slides. The rock samples,
palynological residues and slides are stored in the collection of the Institute of Geological Sciences, Polish Academy of Sciences, Kraków.
RESULTS
All samples contain dinocysts – 121 taxa have been found. Their distribution
and frequency is shown in Figure 4 whereas their alphabetic list is provided in the
Appendix 2. Dinocyst microphotographs are shown in Figures 5–10.
An outstanding feature of dinocyst assemblage from Bêczyn is the frequent occurrence of recycled dinocysts that occur in all samples. Dinocysts, both Miocene
and those treated as recycled, are generally very well preserved. Among the latter,
some specimens of the same species show various states of preservation, from very
well preserved through considerably worse preserved.
The general good preservation of the dinocysts makes it difficult to distinguish
the recycled specimens, especially among species with long stratigraphic ranges including Palaeogene through Early-Middle Miocene. Questionable assignment refers to dinocyst species like Systematophora placacantha, Spiniferites ramosus,
Reticulatosphaera actinocoronata or Palaeocystodinium golzowense, which are
known from Palaeogene and Miocene strata. In this case, specimens showing a
worse state of preservation (especially dark coloured) are tentatively treated as recycled. However, this may be misleading because recycled dinocysts could be
transported into basin coated in host deposit that prevented from mechanical damage – Krach (in Krach & Ksi¹¿kiewicz, 1948, p. 279) mentions frequent occurrence
of tiny rock particles in clay residues evidently derived from the pre-Miocene
substratum.
The Miocene dinocyst assemblage from the Bêczyn clays is dominated by specimens of the genus Batiacasphaera (mainly B. sphaerica), and the species Spiniferites ramosus and Systematophora placacantha. Species like Polysphaeridium zoharyi and Reticulatosphaera actinocoronata occur infrequently throughout the
section (the latter species occur frequently in sample Bnz10). The majority of other
species is found as single or rare specimens in a few samples only (e.g., Cribroperidinium tenuitabulatum, Lingulodinium machaerophorum, Selenopemphix
nephroides, Nematosphaeropsis labyrinthus). Polysphaeridium subtile occurs
mainly in the lower (tentatively – see Materials) part of the section, whereas Pa-
376
Fig. 4.
P. GEDL
Distribution of dinoflagellate cysts in the Middle Miocene clays at Bêczyn
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
Figure 4 continued
377
378
P. GEDL
Figure 4 continued
laeocystodinium striatogranulosum occurs most frequently in the higher part. Impagidinium paradoxum, Impagidinium sp. A and Imperfectodinium septatum occur
in the middle (Bnz9–Bnz11) and uppermost part of the section (Bnz15–Bnz17).
Among the dinocysts treated as recycled, the most frequent are Palaeogene species dominated by the genus Homotryblium. This genus is treated as recycled, although some authors report the occurrence of its representatives in Lower or even
Middle Miocene strata (for discussion see de Verteuil & Norris, 1996). Treating of
the genus Homotryblium as recycled is justified by fact that it has not been found in
other subevaporatic sections of the Miocene sequence in the Polish part of Carpathian Foredeep (Gedl, 1996, 1997). A single specimen of H. pallidum from the
Baranów beds is probably recycled (Gedl, 1999). Other recycled Palaeogene dinocysts include representatives of the subfamily Wetzelielloideae, genus Deflandrea,
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
379
Fig. 5. Dinocysts from the Middle Miocene clays at Bêczyn: in situ taxa. Scale bar (in A)
represents 25 µm and refers to all photomicrographs. Slide code given. A – Achomosphaera sp.,
Bnz17; B – Spiniferites ramosus, Bnz17; C, E – Nematosphaeropsis labyrinthus, C: Bnz16, E: small
specimen, Bnz17; D – Hystrichokolpoma rigaudiae, Bnz9; F–H – Batiacasphaera micropapillata, F:
Bnz17, G: Bnz13, H: Bnz1; I–L – Batiacasphaera sphaerica, I: the same specimen, various foci,
Bnz14, J: Bnz17, K: Bnz12, L: Bnz17; M – Labyrinthodinium truncatum, Bnz3; N – Impagidinium
paradoxum, Bnz17; O – Impagidinium sp. A, Bnz16; P, Q – Imperfectodinium septatum, P: Bnz17,
Q: the same specimen, various foci, Bnz14; R, S – Reticulatosphaera actinocoronata, R: Bnz15, S:
Bnz16; T – Polysphaeridium zoharyi, the same specimen, various foci, Bnz15; U, V – Unipontidinium aquaeductum, U: Bnz17, V: Bnz4; W – Melitasphaeridium choanophorum, the same
specimen, various foci, Bnz1; X, Z – Cribroperidinium tenuitabulatum, X: Bnz8, Z: Bnz15; Y –
Tuberculodinium vancampoe, Bnz1
380
P. GEDL
Fig. 6. Dinocysts from the Middle Miocene clays at Bêczyn: in situ taxa. Scale bar (in A)
represents 25 µm and refers to all photomicrographs. Slide code given. A, B – Selenopemphix
brevispinosa, A: Bnz7, B: Bnz6; C, D – Selenopemphix nephroides, C: Bnz7, D: Bnz16; E, F –
Palaeocystodinium striatogranulosum, E: Bnz17, F: Bnz16; G – Lejeunecysta sp., Bnz4; H –
Dapsilidinium pseudocolligerum, Bnz3; I, J – Operculodinium centrocarpum, I: Bnz17, J: Bnz15;
K–P – Systematophora placacantha, K: the same specimen, various foci, Bnz17, L: Bnz6, M: the
same specimen, various foci, Bnz17, N: Bnz17, O: Bnz17, P: the same small specimen, various foci,
Bnz17
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
381
Fig. 7. Dinocysts from Middle Miocene clays at Bêczyn: recycled Palaeogene taxa. Scale bar (in
B) represents 25 µm and refers to all photomicrographs. Slide code given. A, B – Polysphaeridium
subtile, A: complete specimen, Bnz6, B: Bnz6; C–E – Homotryblium tenuispinosum, C: Bnz2, D:
Bnz2, E: Bnz16; F – Homotryblium aculeatum, Bnz13; G – Homotryblium floripes, Bnz15; H –
Homotryblium abbreviatum, Bnz2; I, J – Tityrosphaeridium cantharellus, I: Bnz16, J: Bnz17; K –
Glaphyrocysta exuberans, Bnz2; L – Cordosphaeridium robustum, Bnz12; M – Cordosphaeridium
funiculatum, Bnz13; N – Areoligera coronata, Bnz12; O – Distatodinium biffi, Bnz12; P – Cordosphaeridium gracile, Bnz17; Q – Thalassiphora pelagica, Bnz16
382
P. GEDL
Fig. 8. Dinocysts from the Middle Miocene clays at Bêczyn: recycled Palaeogene taxa. Scale bar
(in A) represents 25 µm and refers to all photomicrographs. Slide code given. A – Enneadocysta
pectiniformis, Bnz17; B – Gerdiocysta sp., Bnz16; C – Diphyes colligerum, Bnz6; D – Apectodinium
homomorphum, Bnz4; E – Apectodinium parvum, Bnz2; F – Areosphaeridium michoudii, Bnz8; G –
Operculodinium microtriainum, Bnz14; H, J, K – Deflandrea sp., H: Bnz9, J: Bnz9, K: Bnz17; I –
Hystrichokolpoma cinctum, Bnz17; L, M – Wetzeliella sp., L: well preserved specimen, Bnz4, M:
poorly preserved specimen, Bnz4; N – Wetzeliella symmetrica, Bnz15; O – Charlesdowniea sp.,
Bnz11; P – Rhombodinium freienwaldensis, Bnz17; Q – Rhombodinium draco, Bnz7; R – Rhombodinium perforatum, Bnz13; S – Deflandrea oebisfeldensis, Bnz16; T – Cerodinium albertii, Bnz13
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
383
Fig. 9. Dinocysts from the Middle Miocene clays at Bêczyn: recycled Cretaceous taxa. Scale bar
(in L) represents 25 µm and refers to all photomicrographs. Slide code given. A – Achomosphaera
ramulifera, Bnz1; B – Chatangiella sp., Bnz11; C, D – Kleithriasphaeridium eoinodes, both
specimens: Bnz1; E – Impagidinium margaritiferum, Bnz2; F–I – Circulodinium sp., F: Bnz17, G:
Bnz1; H: Bnz1, I: Bnz1; J – ?Cribroperidinium sp., specimen with perforated cyst wall, Bnz1; K –
Cribroperidinium ventriosum, Bnz1; L – Oligosphaeridium complex, Bnz12; M – Cribroperidinium
sp., Bnz1; N, S, U, V – ?Cribroperidinium edwardsii, all specimens: Bnz1, S: isolated operculum; O –
Palaeohystrichophora infusorioides, Bnz6; P–R – Chlamydophorella sp., P: Bnz14, Q: Bnz1, R:
Bnz1; T – ?Oligosphaeridium asterigerum, Bnz1; W – Oligosphaeridium albertense, Bnz1; X –
Oligosphaeridium complex brevispinum, Bnz7
384
P. GEDL
Fig. 10. Dinocysts from the Middle Miocene clays at Bêczyn: recycled Cretaceous and Jurassic
taxa. Scale bar (in K) represents 25 µm and refers to all photomicrographs. Slide code given. A, B –
Callaiosphaeridium asymmetricum, A: Bnz1; B: Bnz12; C – Coronifera oceanica, Bnz1; D –
Dingodinium coerviculum, Bnz15; E – Gonyaulacysta diutina, Bnz2; F – Pterodinium sp., Bnz16; G
– Ovoidinium sp., Bnz1; H, I – Subtilisphaera sp., both specimens: Bnz1; J – Palaeoperidinium
cretaceum, Bnz1; K – Muderongia simplex, the same specimen, various foci: Bnz1; L, Z –
Odontochitina sp., L: Bnz6, Z: Bnz1; M, Q – Odontochitina costata, both specimens: Bnz1, M:
epicyst, Q: hypocyst; N – Pseudoceratium anaphrissum, Bnz1; O – Endoscrinium sp., Bnz2; P –
Valensiella reticulata, Bnz1; R – ?Surculosphaeridium longifurcatum, Bnz1; S – Dinogymnium sp.,
Bnz8; T – Ctenidodinium sp., Bnz1; U – Glossodinium dimorphum, Bnz8; V – Valensiella sp., Bnz1;
W – Systematophora sp., Bnz1; X – Stiphrosphaeridium anthophorum, Bnz1; Y – Kiokansium
polypes, Bnz1
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
385
and several species like Areosphaeridium diktyoplokum, Eatonicysta ursulae,
Glaphyrocysta pastielsii and Diphyes colligerum.
Cretaceous dinocysts are less frequent – they usually occur as single specimens.
These are species like Callaiosphaeridium asymmetricum, ?Cribroperidinium edwardsii, Palaeohystrichophora infusorioides, Isabelidinium sp., Dinogymnium
sp., Odontochitina sp., Dingodinium coerviculum, Oligosphaeridium asterigerum,
Chatangiella sp. The majority of Cretaceous dinocysts from the Bêczyn clays is
typical for Late Cretaceous but some might be recycled from Lower Cretaceous
strata (e.g., ?Cribroperidinium edwardsii, Dingodinium coerviculum, Oligosphaeridium complex, Circulodinium sp., Callaiosphaeridium asymmetricum, Muderongia simplex, Stiphrosphaeridium anthophorum). Stratigraphic ranges of some
Late Cretaceous species extend also into the Tertiary (e.g., Oligosphaeridium complex, Cerodinium albertii, Hystrichosphaeridium sp.).
One specimen of the Middle Jurassic genus Ctenidodinium and one specimen of
the Late Jurassic species Glossodinium dimorphum represent the Jurassic dinocysts
in the Bêczyn clays.
The frequency of recycled dinocyst occurrence (note the difficulties with distinguishing recycled taxa) in the investigated section show some fluctuations, but they
usually represent 30–50% of the dinocyst assemblage (Fig. 11). Recycled dinocysts
are generally most frequent in the lowermost samples: Bnz1, Bnz2 (over 80% of
dinocyst assemblage) and Bnz4 (over 60%). The stratigraphically highest sample,
Bnz17, contains less than 15% of recycled specimens.
BIOSTRATIGRAPHY
The occurrence of Unipontidinium aquaeductum in the investigated clays at
Bêczyn correlates these sediments with interregional dinocyst zone D17 of Costa &
Manum (1988) calibrated to the calcareous nannoplankton zones NN5–NN6 of
Martini (1971). The lowest occurrence of Unipontidinium aquaeductum is an important biostratigraphic event that defines the bases of several dinocyst zones.
These zones are usually calibrated on NN5 and on the lower/lowermost part of zone
NN6 (e.g., Manum, 1976; Costa & Downie, 1979; Piasecki, 1980; Edwards, 1984;
Brown & Downie, 1985; Powell, 1986b; Head et al., 1989; Manum et al., 1989;
Powell, 1992; Zevenboom, 1995).
In Polish part of the Carpathian Foredeep, Unipontidinium aquaeductum has
been recorded from lower Badenian strata only. This species occurs in the Korytnica clays (Gedl, 1996) and the upper part of the Baranów beds (Gedl, unpubl.) –
i.e., the Piñczów Formation sensu Alexandrowicz et al. (1982). In the western part
of the Carpathian Foredeep, this species has been found in the upper part of the
Skawina Formation (Gedl, unpubl.). Thus, the lower Badenian clays from Bêczyn
represent the upper part of the Skawina Formation (sensu Alexandrowicz, 1997).
Its age, based on foraminifera and calcareous nannoplankton, can be calibrated to
the upper part of NN4-lower part of NN6 (Olszewska, 1999; Garecka & Jugowiec,
1999).
386
P. GEDL
Fig. 11. Percentage of in situ and recycled dinocysts in the Middle Miocene clays at Bêczyn
Age-assessment based on dinocysts generally confirms the results of previous
palaeontological studies of Bêczyn clays. Krach (in Krach & Ksi¹¿kiewicz, 1948),
Ma³ecki (1949) and £uczkowska (1957) suggest an early Badenian (early Tortonian) age of the Bêczyn clays.
PALAEOENVIRONMENT
Reconstruction of the palaeoenvironmental setting of the Bêczyn clays based on
dinocysts is impeded due to frequent difficulties in distinguishing recycled dinocysts. However, within the Bêczyn dinocyst assemblage that is treated as Miocene,
dinocysts typical for various environments can be found. The most frequent dino-
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
387
Fig. 12. Palaeogeographic reconstruction of the Carpathian Foredeep basin in the vicinity of
Bêczyn during the early Badenian
cyst genus from Bêczyn, Batiacasphaera sp., has been reported from Miocene
near-shore environments of the Carpathian Foredeep (Gedl, 1996, 1997). Polysphaeridium is another dinocyst genus from Bêczyn, motile stage of which is believed to inhabit near-shore waters (e.g., Wall & Dale, 1969; Dale, 1976; Wall et al.,
1977; Harland, 1983). Different environmental preferences are suggested for the
genus Impagidinium, whose recent representatives frequently occur in offshore waters (e.g., Dale 1996; Rochon et al. 1999; Vink et al. 2000). Offshore preferences
are also suggested for the dinocyst species Unipontidinium aquaeductum and Palaeocystodinium striatogranulosum (Gedl, 1995). The co-occurrence of dinocysts
inhabiting various settings may be explained by hydrodynamic transport responsible for basinward washout of neritic dinocyst species.
Noteworthy, there is no correlation between the frequency of recycled dinocysts
and the frequency of resedimented near-shore specimens. Samples with a high frequency of recycled dinocysts (Bnz2, Bnz4, Bnz8 and Bnz13) contain the lowest
amounts of Batiacasphaera. Moreover, samples with the highest ratio of in situ
dinocysts (Bnz3, Bnz5, Bnz9, Bnz12, Bnz14 and Bnz17) have the highest frequencies of both neritic and offshore species. There is also no correlation between the
amount of resedimented macrofauna and the frequency of recycled dinocysts. This
is clearly visible in the example of samples Bnz13 and Bnz14 (see Fig. 3B). The
former sample taken from clay devoid of macrofauna contains mainly recycled
dinocysts. In contrast, sample Bnz14 taken from a shellbed contain very few preMiocene dinocysts (Fig. 11).
Therefore, it is not certain if the recycled dinocysts have been transported to the
basin during periods of increased erosion and supply of terrigenous material (i.e.,
during floods) or whether they were eroded from submerged pre-Miocene substratum (Fig. 12). Their usual good state of preservation suggests that recycled dinocysts were supplied to the basin coated in rock particles (see Krach & Ksi¹¿kiewicz,
1948, p. 279). Differences in preservation within the same taxon may indicate various source areas during the Middle Miocene. Poorly preserved dinocysts are fre-
388
P. GEDL
quently found in the Carpathian flysch deposits (e.g., Gedl, 2000) whereas excellently preserved forms are known from the epicontinental deposits of the Polish
Lowlands (e.g., Gedl & Ziaja, 2004). The frequent occurrence of well-preserved
pre-Miocene specimens in the clays at Bêczyn suggests that they were derived from
recently non-existing marginal deposits of the Carpathian basin.
CONCLUSIONS
1. The studied Middle Miocene clays at Bêczyn contain frequent and wellpreserved dinocysts. A total of 121 dinocyst taxa have been found.
2. The presence of Unipontidinium aquaeductum indicates that the clays at Bêczyn represent the Unipontidinium aquaeductum Dinocyst Zone, which is correlated
with the NN5–NN6 calcareous nannoplankton Zone.
3. The dinocyst assemblage at Bêczyn consists in large part of recycled, preMiocene specimens. They represent mainly Late Cretaceous and Palaeogene specimens. Their occurrence indicates increased erosion of pre-Miocene substratum in
this area during the early Badenian. It is not certain whether the source area was a
land mass or a submerged ridge built of flysch deposits, or whether it was a rather
recently non-existing margin of the Carpathian basin with epicontinental deposits.
Acknowledgement
I would like to thank Dr B. Ko³odziej (Institute of Geological Sciences, Jagiellonian University,
Kraków) for encouraging me to undertake this study. He is also thanked, together with Dr J. Stolarski
(Institute of Palaeobiology, Polish Academy of Sciences, Warszawa), for assistance during sample
collecting. Dr M. Barski (Faculty of Geology, Warsaw University) is acknowledged for critical
reading the manuscript and Dr M. A. Kaminski is thanked for its linguistic correction.
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DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
391
APPENDIX 1
Location and description of studied samples. Reaction with HCl refers to 38%
acid: +++ – very strong; ++ – strong; + – moderate; = – no reaction
APPENDIX 2
An alphabetical listing of dinocyst taxa found in the Middle Miocene clays at Bêczyn is provided
below. Full taxonomic citations are given in Williams et al. (1998). Taxa considered to be recycled are
identified with an asterisk (*). Numbers in parentheses refer to Figure 4 followed by reference to the
appropriate photomicrographs in Figures 5–10.
Achomosphaera alcicornu (Eisenack 1954) Davey et Williams 1966* (56)
Achomosphaera ramulifera (Deflandre 1937) Evitt 1963* (101; Fig. 9A)
Achomosphaera sp. (27; Fig. 5A)
392
P. GEDL
Adnatosphaeridium vittatum Williams et Downie 1966* (98)
Apectodinium homomorphum (Deflandre et Cookson 1955) Lentin et Williams 1977* (89; Fig. 8D)
Apectodinium parvum (Alberti 1961) Lentin et Williams 1977* (96; Fig. 8E)
Areoligera coronata (O. Wetzel 1933) Lejeune-Carpentier 1938* (60; Fig. 7N)
Areoligera medusettiformis (O. Wetzel 1933) Lejeune-Carpentier 1938* (66)
Areosphaeridium diktyoplokum (Klumpp 1953) Eaton 1971* (38)
Areosphaeridium michoudii Bujak 1994* (45; Fig. 8F)
Batiacasphaera hirsuta Stover 1977 (46)
Batiacasphaera micropapillata Stover 1977 (55; Fig. 5F–H)
Batiacasphaera sphaerica Stover 1977 (1; Fig. 5I–L)
Callaiosphaeridium asymmetricum (Deflandre et Courteville 1939) Davey et Williams 1966* (3; Fig.
10A, B)
Cannosphaeropsis utinensis O. Wetzel 1932* (67)
Cerodinium albertii (Corradini 1973) Lentin et Williams 1987* (57; Fig. 8T)
Charlesdowniea sp.* (16; Fig. 8O)
Chatangiella sp.* (65; Fig. 9B)
Chiropteridium sp.* (63)
Chlamydophorella sp.* (51; Fig. 9P–R)
Chytroeisphaeridia chytroeides (Sarjeant 1962) Downie et Sarjeant 1965* (102)
Circulodinium sp.* (28; Fig. 9F–I)
Cordosphaeridium funiculatum Morgenroth 1966* (53; Fig. 7M)
Cordosphaeridium gracile (Eisenack 1954) Davey et Williams 1966* (20; Fig. 7P)
Cordosphaeridium inodes (Klumpp 1953) Eisenack 1963* (59)
Cordosphaeridium robustum (Gocht 1969) Sarjeant 1981* (61; Fig. 7L)
Coronifera oceanica Cookson et Eisenack 1958* (103; Fig. 10C)
Corrudinium sp.* (71)
?Cribroperidinium edwardsii (Cookson et Eisenack 1958) Davey 1969* (35; Fig. 9N, S, U, V)
Cribroperidinium tenuitabulatum (Gerlach 1961) Helenes 1984 (43; Fig. 5X, Z)
Cribroperidinium ventriosum (O. Wetzel 1933) Lentin et Williams 1973* (104; Fig. 9K)
Cribroperidinium sp.* (105; Fig. 9K, M)
?Cribroperidinium sp.* (106; Fig. 9J)
Ctenidodinium sp.* (107; Fig. 10T)
Dapsilidinium pseudocolligerum (Stover 1977) Bujak, Downie, Eaton et Williams 1980 (91; Fig. 6H)
Deflandrea heterophlycta Deflandre et Cookson 1955* (69)
Deflandrea oebisfeldensis Alberti 1959* (33; Fig. 8S)
Deflandrea sp.* (10; Fig. 8H, J, K)
Dingodinium cerviculum Cookson et Eisenack 1958* (40; Fig. 10D)
Dinogymnium sp.* (79; Fig. 10S)
Diphyes colligerum (Deflandre et Cookson 1955) Cookson 1965* (86; Fig. 8C)
Distatodinium biffi Brinkhuis, Powell et Zevenboom 1992* (62: Fig. 7O)
Eatonicysta ursulae (Morgenroth 1966) Stover et Evitt 1978* (90)
Endoscrinium sp.* (97; Fig. 10O)
Enneadocysta pectiniformis (Gerlach 1961) Stover et Williams 1995* (11; Fig. 8A)
Exochosphaeridium sp.* (75)
Gerdiocysta sp.* (31; Fig. 8B)
Glaphyrocysta exuberans (Deflandre et Cookson 1955) Stover et Evitt 1978* (99; Fig. 7K)
Glaphyrocysta pastielsii (Deflandre et Cookson 1955) Stover et Evitt 1978* (93)
Glaphyrocysta semitecta (Bujak 1980) Lentin et Williams 1981* (78)
Glaphyrocysta sp.* (29)
Glossodinium dimorphum Ioannides, Stavrinos et Downie1977* (77; Fig. 10U)
Gonyaulacysta diutina Duxbury 1977* (95; Fig. 10E)
Homotryblium abbreviatum Eaton 1976* (41; Fig. 7H)
Homotryblium aculeatum Williams 1978* (58; Fig. 7F)
DINOCYSTS FROM MIDDLE MIOCENE OF CARPATHIAN FOREDEEP
393
Homotryblium floripes ((Deflandre et Cookson 1955) Stover 1975* (52; Fig. 7G)
Homotryblium pallidum Davey et Williams 1966* (83)
Homotryblium plectilum Drugg et Loeblich 1967* (24)
Homotryblium tenuispinosum Davey et Williams 1966* (22; Fig. 7C–E)
Homotryblium sp.* (6)
Hystrichokolpoma cinctum Klumpp 1953* (36; Fig. 8I)
Hystrichokolpoma rigaudiae Deflandre et Cookson 1955 (74; Fig. 5D)
Hystrichosphaeridium sp.* (7)
Impagidinium brevisulcatum Michoux 1985* (82)
Impagidinium cristatum (May 1980) Lentin et Williams 1981 (39)
Impagidinium margaritiferum (Cookson et Eisenack 1960) Stover et Evitt 1978* (94; Fig. 9E)
Impagidinium paradoxum (Wall, 1967) Stover et Evitt 1978 (15; Fig. 5N)
Impagidinium sp. A (14; Fig. 5O)
Impagidinium sp. (87)
Imperfectodinium septatum Zevenboom et Santarelli 1995 (18; Fig. 5P, Q)
Isabelidinium sp.* (12)
Kiokansium polypes (Cookson et Eisenack 1962) Below 1982* (108; Fig. 10Y)
Kleithriasphaeridium eoinodes (Eisenack 1958) Davey 1974* (109; Fig. 9C, D)
Labyrinthodinium truncatum Piasecki 1980 (92; Fig. 5M)
Lejeunecysta sp. (38; Fig. 6G)
Lingulodinium machaerophorum (Deflandre et Cookson 1955) Wall 1967 (47)
Melitasphaeridium choanophorum (Deflandre et Cookson 1955) Harland et Hill 1979 (23; Fig. 5W)
Muderongia simplex Alberti 1961* (110; Fig. 10K)
Nematosphaeropsis labyrinthus (Ostenfeld 1903) Reid 1974* (13; Fig. 5C, E)
Odontochitina costata Alberti 1961* (111; Fig. 10M, Q)
Odontochitina sp.* (76; Fig. 10L, Z)
Oligosphaeridium albertense (Pocock 1962) Davey et Williams 1969* (112; Fig. 9W)
?Oligosphaeridium asterigerum (Gocht 1959) Davey et Williams 1969* (72; Fig. 9T)
Oligosphaeridium complex (White 1842) Davey et Williams 1966* (64; Fig. 9L)
Oligosphaeridium complex brevispinum Jain 1977* (80; Fig. 9X)
Operculodinium centrocarpum (Deflandre et Cookson 1955) Wall 1967 (8; Fig. 6I, J)
Operculodinium microtriainum (Klumpp 1953) Islam 1983* (50; Fig. 8G)
Ovoidinium sp.* (113; Fig. 10G)
Palaeocystodinium golzowense Alberti 1961 (19)
Palaeocystodinium striatogranulosum Zevenboom et Santarelli 1995 (9; Fig. 2E, F)
Palaeohystrichophora infusorioides Deflandre 1935* (84; Fig. 9O)
Palaeoperidinium cretaceum Pocock 1962* (114; Fig. 10J)
Polysphaeridium subtile Davey et Williams 1966 (85; Fig. 7A, B)
Polysphaeridium zoharyi (Rossignol 1962) Bujak, Downie, Eaton et Williams 1980 (17; Fig. 5T)
Pseudoceratium anaphrissum (Sarjeant 1966) Bint 1986* (115; Fig. 10N)
Pterodinium sp.* (37; Fig. 10F)
Reticulatosphaera actinocoronata (Benedek 1972) Bujak et Matsuoka 1986 (32; Fig. 5R, S)
Rhombodinium draco Gocht 1955* (81; Fig. 8Q)
Rhombodinium freienwaldensis (Gocht 1955) Costa et Downie 1979* (2; Fig. 8P)
Rhombodinium perforatum (Jan du Chêne et Châteauneuf 1975) Lentin et Williams 1977* (54; Fig.
8R)
Rhombodinium sp. sensu Gedl 2000* (68)
Selenopemphix brevispinosa Head, Norris et Mudie 1989 (73; Fig. 6A, B)
Selenopemphix nephroides Benedek 1972 (34; Fig. 6C, D)
Spiniferites pseudofurcatus (Klumpp 1953) Sarjeant 1970 (49)
Spiniferites ramosus (Ehrenberg 1838) Loeblich et Loeblich 1966 (4; Fig. 5B)
Stiphrosphaeridium anthophorum (Cookson et Eisenack 1958) Lentin et Williams 1985* (116; Fig.
10X)
394
P. GEDL
Subtilisphaera sp.* (117; Fig. 10H, I)
?Surculosphaeridium longifurcatum (Firtion 1952) Davey, Downie, Sarjeant et Williams 1966* (118;
Fig. 10R)
Surculosphaeridium sp.* (70)
Systematophora placacantha (Deflandre et Cookson 1955) Davey, Downie, Sarjeant et Williams
1966 (5; Fig. 6K–P)
Systematophora sp.* (119; Fig. 10W)
Tectatodinium sp. (48)
Thalassiphora pelagica (Eisenack 1954) Eisenack et Gocht 1960* (30; Fig. 7Q)
Tityrosphaeridium cantharellus (Brosius 1963) Sarjeant 1981* (26; Fig. 7I, J)
Tuberculodinium vancampoe (Rossignol 1962) Wall 1967 (100; Fig. 5Y)
Unipontidinium aquaeductum (Piasecki 1980) Wrenn 1988 (21; Fig. 5U, V)
Valensiella reticulata (Davey 1969) Courtinat 1989* (120; Fig. 10P)
Valensiella sp.* (121; Fig. 10V)
Wetzeliella symmetrica Weiler 1956* (44; Fig. 8N)
Wetzeliella sp.* (25; Fig. 8L, M)
“round-browns” (42)

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