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STUDIA GEOLOGICA POLONICA Vol. 124, Kraków 2005, pp. 199–214. Methods and Applications in Micropalaeontology Edited by J. Tyszka, M. Oliwkiewicz-Miklasiñska, P. Gedl & M. A. Kaminski Andrzej SZYD£O 1 Benthic foraminiferal morphogroups and taphonomy of the Cieszyn beds (Tithonian–Neocomian, Polish Outer Carpathians) (Figs 1–4) Abstract. Test gross morphology and taphonomy of calcareous benthic foraminifers from the Cieszyn beds were mainly controlled by depositional parameters, paleoecological conditions and paleogeographical settings. All of these control factors were influenced by a major tectonic event represented by the rifting of the Cieszyn Basin. In its earliest history it acted as a gulf adjacent to the Eastern European Platform. The Tithonian marls and marly shales known in Polish regional geology as the lower Cieszyn shales and Cieszyn limestones yielded highly diverse calcareous benthic foraminiferal assemblages which are dominated by shallow and deep infaunal morphogroups. A remarkably well-developed epifaunal component can be recognized as well. Shelf and peri-reefal foraminifera originally developed their life cycles under well-oxygenated conditions in the immediate proximity of the sea-bottom. The initiation of turbidite sedimentation lead to the removal and transportation of shelf foraminifers into deeper water environments (bathyal). Consequently, the tests of calcareous foraminifers are small, and poorly preserved, presenting dissolution traces, and abraded and corroded surfaces. The mixed and reworked microfauna were replaced by a more uniform one in the proximity of the Jurassic/Cretaceous boundary. Active omnivorous infauna survived not only the change of deposition system but also the deepening of the Cieszyn Basin during Early Cretaceous times. Periodically the supply of reworked and allochthonous calcareous benthic forms into the Cieszyn Basin took place during the Berriasian and Late Valanginian–Hauterivian times. Key words: palaeoecology, benthic foraminifera, morphogroup analysis, Jurassic, Cretaceous, Carpathians, Silesian Unit. 1 Carpathian Branch, Polish Geological Institute, ul. Skrzatów 1, 31-560 Kraków, Poland. E-mail: [email protected]

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Page 1: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

S T U D I A G E O L O G I C A P O L O N I C AVol. 124, Kraków 2005, pp. 199–214.

Methods and Applications in MicropalaeontologyEdited by J. Tyszka, M. Oliwkiewicz-Miklasiñska,

P. Gedl & M. A. Kaminski

Andrzej SZYD£O1

Benthic foraminiferal morphogroups and taphonomy

of the Cieszyn beds (Tithonian–Neocomian,

Polish Outer Carpathians)

(Figs 1–4)

Abstract. Test gross morphology and taphonomy of calcareous benthic foraminifers from theCieszyn beds were mainly controlled by depositional parameters, paleoecological conditions andpaleogeographical settings. All of these control factors were influenced by a major tectonic eventrepresented by the rifting of the Cieszyn Basin. In its earliest history it acted as a gulf adjacent to theEastern European Platform. The Tithonian marls and marly shales known in Polish regional geologyas the lower Cieszyn shales and Cieszyn limestones yielded highly diverse calcareous benthicforaminiferal assemblages which are dominated by shallow and deep infaunal morphogroups. Aremarkably well-developed epifaunal component can be recognized as well. Shelf and peri-reefalforaminifera originally developed their life cycles under well-oxygenated conditions in the immediateproximity of the sea-bottom. The initiation of turbidite sedimentation lead to the removal andtransportation of shelf foraminifers into deeper water environments (bathyal). Consequently, the testsof calcareous foraminifers are small, and poorly preserved, presenting dissolution traces, and abradedand corroded surfaces. The mixed and reworked microfauna were replaced by a more uniform one inthe proximity of the Jurassic/Cretaceous boundary. Active omnivorous infauna survived not only thechange of deposition system but also the deepening of the Cieszyn Basin during Early Cretaceoustimes. Periodically the supply of reworked and allochthonous calcareous benthic forms into theCieszyn Basin took place during the Berriasian and Late Valanginian–Hauterivian times.

Key words: palaeoecology, benthic foraminifera, morphogroup analysis, Jurassic, Cretaceous,Carpathians, Silesian Unit.

1 Carpathian Branch, Polish Geological Institute, ul. Skrzatów 1, 31-560 Kraków, Poland. E-mail:[email protected]

Page 2: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

INTRODUCTION

The uppermost Jurassic and Neocomian sediments in the Polish Outer Carpathi-ans known as the Cieszyn beds, are more completely developed in the Silesian Unitof the Cieszyn Foothills (Fig. 1). These sediments were deposited in the CieszynBasin, which was the initial basin in the Polish part of the Carpathians. During thedeposition of the Cieszyn beds this basin was transformed from a marginal, narrowbasin close to a carbonate platform into a deep marine basin on the northern marginsof the Tethys. At the time, the formation of the Cieszyn Basin had a fundamental in-fluence on the evolution of different biotopes, which were colonized by foramini-feral population including calcareous taxa.

Foraminifers from the studied deposits were firstly described by Geroch (1959;1960; 1961). His studies as well as most of subsequent ones carried out by Bieda etal. (1963), Geroch & Nowak (1963), Geroch (1966), Geroch et al. (1967), Geroch& Olszewska (1990), Kaminski et al. (1992), Geroch & Kaminski (1995) focusedon the agglutinated foraminiferal assemblages. Biostratigraphical zonation of theCieszyn beds is also based on foraminiferal taxa of this group (Geroch & Nowak,1984; Olszewska, 1997) (Fig. 2). Calcareous benthic taxa were occasionally re-ported (Geroch, 1966; Bielecka & Geroch, 1977; Olszewska, 1984). This group ofmicrofossils was extensively analyzed by Szyd³o (1997) and Szyd³o & Jugowiec(1999). Additional data concerning the morphogroups and taphonomy of aggluti-nated foraminifers from the Cieszyn beds were published by Szyd³o (2002a; 2004).All these studies provided the opportunity to compare morphological and tapho-nomic trends in both groups of foraminifera, calcareous and agglutinantsrespectively.

200 A. SZYD£O

Fig. 1. The distribution of the Cieszyn beds and studied profiles in the Cieszyn Foothills. 1–4Cieszyn subunit and Subsilesian Unit: 1 – lower Cieszyn shales, 2 – Cieszyn limestones, 3 – upperCieszyn shales, 4 – other undivided deposits; 5 – Godula subunit, 6 – line of overthrust. FromKsi¹¿kiewicz (1964) and Nowak (1973), modified

Page 3: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

GEOLOGICAL BACKGROUND

The present study is based on samples from the Cieszyn beds. Tectonically,these deposits belong to the Cieszyn tectonostructural subunit, which is includedwithin the Silesian Unit (Fig. 1). Traditionally, the Cieszyn beds are divided into thelower Cieszyn shales, the Cieszyn limestones and the upper Cieszyn shales (Biedaet al., 1963) (Fig. 2).

The lower Cieszyn shales (Tithonian in age) consist mainly of marls and marlyshales, and are subdivided into three informal units known from the base to the topas (1) productive non-flysch complex (lower part), (2) rhythmic sequence with car-bonates and (3) partly reworked series containing carbonate olistolites and exotics.The Cieszyn limestones (Upper Tithonian–Berriasian) are a carbonate complexcomprised layered marls and marly shales. Limey shales from the upper Cieszynshales (Valanginian–Hauterivian) are interbedded with carbonates and by sand-stones in the lowermost and upper parts of the unit respectively (Fig. 3).

APPROACH TO METHODS

The samples were collected from outcrops of the Cieszyn beds, which occur inthe Cieszyn Foothills. The studied profiles are located near Cieszyn (Olza, Puñców,Leszna Górna, Cisownica, Gumna, Ogrodzona, Goleszów-Marglownia), Skoczów

BENTHIC FORAMINIFERAL MORPHOGROUPS 201

Fig. 2. Lithology and stratigraphical position of the Cieszyn beds

Page 4: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

(Grodziec-Górki Wielkie, Rudów) and Bielsko-Bia³a (Jasienica, Kamienica,Lipnik) (Figs 1, 3). Foraminifers were sampled from marls, marly shales and limyshales, which were disintegrated by boiling and freezing.

Calcareous foraminifers found in the washed residues represent the data sourcefor the present morphogroups and taphonomic analyses. The 63 µm fraction wasused for the foraminiferal analysis. The microfauna was described with special fo-cus on the relationship between test morphology and living or feeding strategy ofRecent foraminifers (Severin, 1983; Bernhard, 1986; Koutsoukos et al., 1990).Taphonomic studies concern morphogroup affiliation, foraminiferal abundance,size of specimens, test preservation, distribution and accumulation of test in thesediments (Renzi et al., 2002). Taphonomic features were analyzed in foraminif-eral assemblages that typically include 50 to 60, and rarely 100 specimens amongwhich calcareous forms were dominant and diverse (in the lower Cieszyn shalesand the Upper Tithonian Cieszyn limestones) or rarely (in the Berriasian Cieszynlimestones and the upper Cieszyn shales) component. The results were consideredin a broader sense, namely related to changes in depositional patterns and basintectonics.

202 A. SZYD£O

Fig. 3. Studied and sampled profiles of the Cieszyn beds (for lithological setting see Fig. 2) (afterSzyd³o, 2002, modified)

Page 5: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

BENTHIC FORAMINIFERAL MORPHOGROUPS

Calcareous benthic foraminifers from the Cieszyn beds are assigned to six mor-phogroups (Fig. 4). The most of them were already illustrated (Szyd³o, 1997;Szyd³o & Jugowiec, 1999) or have been recently documented (Szyd³o, in press).

Morphogroup 1 (M1) comprises mainly discoidal, flattened or conical testsconsisting of two chambers. They belong to the genera: Andersenolina, Trocholina,Neotrocholina, Ichnusella and Sprillina or Planispirillina. Forms with flat tests(e.g., Trocholina solecensis, Neotrocholina molesta), convex tests (e.g., Ander-senolina elongata, Trocholina involuta), and slightly concave umbilical side (e.g.,Patellina feifeli) belong also to morphogroup 1. In the lower Cieszyn shales (pro-files: Olza, Puñców, Goleszów-Marglownia, Cisownica, Gumna) this morpho-group is dominated by representatives of the family Involutiniidae – (e.g., T. sole-censis, Neotrocholina molesta, Andersenolina alpina, A. elongata) and to them rarePlacentulinidae (e.g., Paalzowella feifeli), and Spirillinidae are added (see AP-PENDIX). Similar morphotypes also occur in the Upper Tithonian Cieszyn lime-stones (profiles: Olza, Puñców, Cisownica, Górki Wielkie-Grodziec). Abundantforaminiferal assemblages consisting mainly of Neotrocholina (e.g., N. molesta)are commonly associated with minor taxa such as Ichnusella burlini (Gorbachik)and Spirillina ssp. Flattened planispiral forms such as Patellina feifeli (Paalzow),Spirillina infima Strickland and Planispirillina flava (Sztejn) (see APPENDIX)were recorded in the upper Cieszyn shales (profiles: Puñców, Cisownica) (Fig. 4).Foraminifers with conical-shape test belonging to the trocholinid group preferperi-reefal environments (Arnaud-Vanneau et al., 1988; Neagu, 1995). Planispiralforms occur in quite diverse paleoecological niches, taxa with ornamented tests be-ing able to live in low-oxygen environments (Bernhard, 1986). Recent epifaunal fo-raminifers belong to active deposit feeders (Koutsoukos et al. 1990).

Morphogroup 2 (M2) consists mainly of flattened-convex and trochospiraltaxa with many chambers and a raised spiral side (e.g., Gavelinella bettenstaedti) towhich those having slightly concave umbilical area: (e.g., Discorbis crimicus, Con-orbina heteromorpha, Reinholdella hofkeri) can be added (see APPENDIX).

These foraminifers are herbivorous epifauna and they dominate in the shelf bio-tope (Koutsoukos et al. 1990). The Morphogroup 2 is overall represented by quite afew forms, which occur at certain stratigraphical levels in the Cieszyn limestones(Goleszów profile) and the upper Cieszyn shales (Puñców profile) (Fig. 4).

Morphogroup 3 (M3) comprises spherical or ovate multichambered taxa, mostof them presenting an elongated test. This morphotype is represented mostly by:Eoguttulina (e.g., E. liassica, E. polygona) and Guttulina (e.g., G. multistriata) (seeAPPENDIX). Unilocular forms belonging to Lagena apiculata neocomiensisBartenstein & Brand were included here by analogy with agglutinant taxa of themorphogroup 2-a (Nagy et al., 1995). The above-mentioned foraminifers aremainly deposit feeders, a group of taxa which develop their life-cycles in the prox-imity of the water/sediment interface (epifauna/shallow infauna). These rather gen-eralized forms feed on plants and bacteria. Detritivore and omnivorous infauna is

BENTHIC FORAMINIFERAL MORPHOGROUPS 203

Page 6: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

204 A. SZYD£O

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Page 7: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

also a component of this morphogroup (Koutsoukos et al. 1990). This morpho-group is frequently encountered in the lower Cieszyn shales (profiles: Olza,Goleszów-Marglownia, Cisownica, Gumna). It occurs at certain stratigraphicallevels in the Upper Tithonian Cieszyn limestones (Fig. 4).

Morphogroup 4 (M4) is less diverse than the previous ones, being representedby only a few taxa. Elongated, cylindrical, and uniserial forms with a straight toslightly curved test and circular section belong to this morphogroup. Uniserialforms with a smooth test presenting chambers of various shape ranging from oval(e.g., Laevidentalina communis, L. debilis, Pseudonodosaria humilis) to angular(e.g., Tristix acutangula, T. explanata, T. temirica, and T. somaliensis) were recog-nized (see APPENDIX). They are omnivorous taxa living close to sea-bottom(Koutsoukos et al. 1990). Occurrences of taxa assigned now to this morphotype arecommon in the lower Cieszyn shales and are reduced to single specimen-occurrences in the Cieszyn Limestones (e.g., Laevidentalina) and the upper Ci-eszyn shales (e.g., Pseudonodosaria) (Fig. 4). The above described morphotypeswere noted in the following profiles: Olza, Puñców, Ogrodzona, Œwiêtoszówka.

Morphogroup 5 (M5) includes mainly elongated forms with flattened symmet-rical and slightly asymmetrical tests (e.g., Ichthyolaria nikitini). This morphogroupis highly diverse including a variety of taxa among which lanceolate and finny(Frondicularia inderica and Citharinella pomeraniae, C. portlandensis respec-tively) tests, both flattened (e.g., Geinitzinita kcyniensis, G. wolinensis) or orna-mented: (e.g., Citharina brevis, C. paucicostata, C. raricostata and C. virgatis) areregarded now as characteristics of this morphogroup (see APPENDIX). Most of theplanispiral taxa displaying rounded margins and flattened tests are included here(e.g., Planularia cordiformis, P. multicostata, Astacolus calliopsis, A. eritheles, A.primus franconicus etc.). This morphogroup is even more diverse, a variety of otherforms being included here (e.g., Marginulinopsis bettenstaedti, M. buskensis, M.robusta, M. striatocostata, Marginulina declivis, M. inaequalis, M. schlönbachi,Vaginulinopsis embaensis) (see APPENDIX). A special remark is made now forthe elongated and planispiral tests showing a sharp margin in contrast to all the otherspecies included here (e.g., Palmula crepidularis, Saracenaria alata-angularis, S.pravoslavlevi) (APPENDIX, Fig. 4). The above mentioned morphological group isknown from shelf areas and belongs to active omnivorous infauna, which dwell atthe water-sediment interface (Koutsoukos et al., 1990). Component taxa of thismorphogroup are also polarized with respect to their paleoecological preferences:on one hand forms with straight and non ornamented tests which prefer environ-ments with normal values of dissolved oxygen (e.g., Lingulina loryi, Planulariapoljenovae, P. subhumilis, P. uilensis) (see APPENDIX) and taxa with ornamentedtest with affinities for low oxygen environments (Bernhard, 1986) on the other.Morphogroup 5 is frequent in the lower Cieszyn shales (profiles: Puñców, Go-leszów-Marglownia, Gumna, Cisownica,) and occasionally encountered in the Up-per Tithonian Cieszyn limestones (Goleszów-Marglownia profile). It only occursin a single profile of the Berriasian Cieszyn limestones (Górki Wielkie-Grodziecprofile) and upper Cieszyn shales (Puñców profile) respectively.

BENTHIC FORAMINIFERAL MORPHOGROUPS 205

Page 8: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

Morphogroup 6 (M6) comprises biconvex forms, mostly lagenids, whichdominate the foraminiferal assemblages in the Cieszyn beds (e.g., Lenticulina am-banjabensis, L. dogieli, L. infravolgaensis, L. muensteri, L. nodosa L. ponderosa,L. quenstedti, L. varians, L. vistulae elongata). Lamarckina (e.g., L. asteriaformis)and Pseudolamarckina (e.g., P. polonica and P. reussi) are among the minor taxaincluded in this morphogroup (APPENDIX, Fig. 4). Similar modern mobile om-nivorous forms belong to epifauna and deep infauna. Diverse Lenticulina assem-blages were considered indicative for environments with high levels of dissolvedoxygen by Bernhard (1986) and Koutsoukos et al. (1990). Morphogroup 6 is bothmost diverse and rich in the lower Cieszyn shales, being often limited to single ge-nus occurrences in the Cieszyn limestones and upper Cieszyn shales (Fig. 4). Theabove described foraminifers were noted in the following profiles: Olza, Puñców,Goleszów-Marglownia, Cisownica, Gumna-Ogrodzona, Œwiêtoszówka, Jasie-nica-Jaworze, Kamienica).

TAPHONOMIC REMARKS AND ASPECTS OF BENTHIC

FORAMINIFERAL ASSEMBLAGE FLUCTUATIONS RELATED TO

SEDIMENTATION EPISODES

Calcareous benthic foraminifers are most frequent and diverse in the lower Ci-eszyn shales. The distribution of the microfauna in these deposits is highly variable.The scarce assemblages recorded in the slumped and slide sediments of the lowerpart of the stratigraphical column gradually change into assemblages consisting ofvaried and common calcareous forms in the middle hemipelagic part of the lowerCieszyn shales. In the uppermost part of the lower Cieszyn shales (carbonate olisto-lites and exotics) calcareous foraminifers are represented only by poor assem-blages. The calcareous benthos was represented by shallow and deep infauna (M5),epifauna (M1), mobile infauna (M6) and also by the very shallow infauna (M3).The epifauna (M1) dominate in the youngest part of the lower Cieszyn shales. Cal-careous benthics were commonly accompanied by forms which agglutinate car-bonate particles and represent active infaunal morphotypes (Szyd³o, 2004).

Generally, the above-mentioned forms are of similar sizes. Those from olis-tostromes are larger. The whole calcareous microfauna are rather poorly preservedand their tests are usually corroded.

The Upper Tithonian Cieszyn limestones yield locally similar but also less di-verse assemblages (nearly 10 genera), while the whole complex contains distinctlyless numerous calcareous foraminifers, which belong usually to 1–3 genera (mainlyrepresentatives of Lenticulina). Episodically more diverse calcareous foraminifers(4 or 5 genera) occur in the Berriasian Cieszyn limestones. These forms belong toactive omnivorous feeders (M1, M5). Most of the calcareous microfauna from theCieszyn limestones are badly preserved. Mobile (M6) and shallow infauna (M4)are dominant. Sometimes herbivorous epifauna (M2) occur among this group offoraminifera (Fig. 4). Monogenic assemblages of arenaceous foraminifers belong-

206 A. SZYD£O

Page 9: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

ing to tubular epifaunal, elongated infaunal and globular surficial, semi-infaunalmorphological groups largely dominate in these deposits (Szyd³o, 2004).

Rare calcareous benthic forms belonging to mobile (M6) and shallow infauna(M4) (Lenticulina, Pseudonodosaria) are usually observed in the upper Cieszynshales (Fig. 4). These forms co-occur with impoverished agglutinant assemblagestypical of deeper waters. Additionally, epifaunal taxa belonging to Sprillinidae andPlanispirillinidae (M1) are observed in the upper part of this unit which also yieldsdeep infaunal forms presenting agglutinated calcareous cement (Szyd³o, 2004).

DISCUSSION

The diversity of calcareous benthic forms is directly controlled by palaeoeco-logical factors, which were determined mainly by the evolution of sedimentary re-gimes, and palaeogeographical settings in the Cieszyn Basin. The earlier Tithonianforaminiferal biocoenosis occurs on a wide shelf. High diversity and abundance ofthese assemblages reflect aerobic bottom conditions favoring morphotypes withvariable test morphology (Koutsoukos et al., 1990). The predominance of infaunaamong these foraminifers can be correlated with an increased food supply (Corliss& Chen, 1988; Corliss & Fois, 1991). Geotectonic reorganization of the Jurassicshelves resulted in environmental instability which is apparent in the small sizes ofthe foraminiferal taxa. Areas with shallow-water carbonate sedimentation werestrongly rebuilt during Tithonian times (Nowak, 1975; S³omka, 1986). Most likelythese wide shelves were rising and subsiding at various rates during that time. Fora-minifers were the subject of removal and transportation into deeper parts of the ba-sin. Their tests document severe dissolution, presenting often abraded and corrodedsurfaces as a direct result of the two processes.

Highly diverse infaunal groups dominated mainly in the lower Cieszyn shalesand occurred locally in the Cieszyn limestones (Fig. 4). The abundance and vari-ability of these foraminifers strongly decreases in the Late Tithonian. This change isregarded now as a direct response of the foraminiferal populations to the replace-ment of non-flysch sedimentation by turbidite deposition. The abrupt changes inthe sedimentary regime over wide shelf areas was followed by strong erosion(Ksi¹¿kiewicz, 1971) and resulted in removal and transportation of the peri-reefaland shelf foraminifers down slope into the Cieszyn Basin. Then, epifauna belong-ing to both active deposit feeders (M1) and herbivore epifauna (M2) occurred incarbonates breccias and carbonate deposits with olistolites and exotics. After thisshort episode paleoecological conditions in the Cieszyn Basin drastically changed.Opportunistic and active infauna belonging mainly to the genus Lenticulina (M6)and sometimes to Pseudonodosaria and Laevidentalina (M4) were usually noted inthe turbidite deposits (the Cieszyn limestones and the upper Cieszyn shales) (Fig.4). The low diversity and decrease in abundance of calcareous foraminifers whichaccompanied almost monogenic agglutinated assemblages belonging to omnivo-rous deposit feeders were related to the establishment of dysaerobic bottom condi-tions in the Early Cretaceous Cieszyn Basin (Bernhard, 1986; Koutsoukos et al.,

BENTHIC FORAMINIFERAL MORPHOGROUPS 207

Page 10: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

1990; Szyd³o, 2004). At that time, the CCD level and a lysocline were raised signifi-cantly and low calcium carbonate levels were noted in the basin. Palaeoenviron-mental conditions in the stratified and deep basin (S³omka, 1986; Malik, 1986)were extremely hostile to life. More frequent transportation of calcareous foramini-fers took place during Berriasian and in Valanginian–Hauterivian.

CONCLUSIONS

Cieszyn Beds yielded mainly reworked and transported calcareous benthic fora-minifers. Most of the microfaunal elements were displaced into bathyal environ-ments during turbiditic episodes and olistolite emplacements in Tithonian times.Foraminifers were also transported into the basin by suspension currents. Periodi-cally, calcareous forms developed abundant assemblages during the Berriasian andLate Valanginian–Hauterivian. Longtime mechanical transport from the shelf en-vironments resulted in the poor preservation of the locally abundant foraminiferalcalcareous tests. Small-sized calcareous forms lived in the area with shallow-watercarbonate sedimentation, the latter being sensitive to the relative sea-level changes.In the Tithonian the microfauna existed under aerobic bottom conditions, which fa-vored the development of different morphotypes. The disappearance from the rec-ord of mixed morphogroups and the occurrence of selected morphotypes belongingto omnivorous feeders corresponds to a change in the paleogeographical and pa-leoecological regimes in the basin in the proximity of the Jurassic/Cretaceousboundary. Dysaerobic conditions in the deepening basin resulted in the develop-ment of rather homogenous and impoverished calcareous foraminiferal assem-blages. Probably the supply of more numerous calcareous forms into the Early Cre-taceous basin was associated with sea-level fluctuations and mostly with the de-struction and erosion of shelf areas at that time.

Acknowlegements

I thank Drs M. Dan Georgescu (University of Saskatchewan) and Jaros³aw Tyszka (PolishAcademy of Science) for their constructive comments on the manuscript and Teresa Mrozek (PolishGeological Institute) for reviewing this paper. I am grateful to Dr Piotr Nescieruk (Polish GeologicalInstitute) for his help during field work and the geological study. The research was partly financialsupported by the State Committee for Sciences Research (6 P04D 041 17).

APPENDIX

The following list includes calcareous species recognized in this study. The above-presentedmorphogroup affiliation and the taxonomical classification proposed by Loeblich and Tappan (1987)and updated by Meyn and Vespermann (1994) were used.

Morphogroup 1

Andersenolina alpina (Coscinoconus alpinus Leupold, Leupold & Bigler,1935: 610, pl. 18, figs 1–11)Andersenolina elongata (Coscinoconus elongatus Leupold, Leupold & Bigler, 1935: 617, pl. 8, figs12–14)Trocholina involuta Mantsurova (T. involuta Mantsurova, Mantsurova & Gorbachik, 1982: 127, pl. 3,fig. 7, pl. 4, fig. 3)

208 A. SZYD£O

Page 11: Benthic foraminiferal morphogroups and taphonomy of the Cieszyn

Trocholina solecensis (T. solecensis Bielecka & Po¿aryski, 1954: 69, pl. 11, fig. 57a–c)Neotrocholina molesta (Trocholina molesta Gorbachik, 1959: 79, pl. 4, figs 1, 2)Ichnusella burlini (Trocholina burlini Gorbachik, 1959: 81, pl. 4, figs 3–5)Spirillina infima (Orbis infimus Strickland, 1846: 30, fig. a)Spirillina polygyrata (S. polygyrata Gümbel, 1862: 214, pl.4, fig. 11a–c)Planispirillina flava (Spirillina flava Sztejn, 1957: 77, pl. 9, fig. 87)Patellina feifeli (Trocholina feifeli Paalzow, 1932: 140, pl. 2, figs 6, 7)Paalzowella feifeli (Trocholina feifeli Paalzow, 1932: 140, pl. 11, figs 4, 6, 7)

Morphogroup 2

Reinholdella hofkeri (Conorbis hofkeri Bartenstein & Brand, 1951: 325, 326, pl. 11, fig. 320)Conorbina heteromorpha (C. heteromorpha Gorbachik, 1971: 134, pl. 5, fig. 7)Discorbis crimicus (D. crimicus Schokhina; Gorbachik & Schokhina, 1960: 102, 12, fig. 1)Gavelinella bettenstaedti (G. bettenstaedti Dieni & Massari, 1966: 175, pl. 8, figs 17a–19b)

Morphogroup 3

Lagena apiculata neocomiensis (L. apiculata neocomiensis Bartenstein & Brand, 1951: 317, pl. 10,figs 275, 276, pl. 13, figs 44–47, 49, 51, pl. 19, figs 43–45)Eoguttulina liassica (Polymorphina liassica Strickland, 1846: 30, fig. b)Eoguttulina polygona (Polymorphina polygona Terquem, 1864 : 305, pl. 14, figs 16–21, 35)Guttulina multistriata (G. multistriata Bielecka, 1975: 353–354, pl.10, figs 14–16)

Morphogroup 4

Laevidentalina communis (Nodosaria communis d’Orbigny, 1826: 254)Laevidentalina debilis (Marginulina debilis Berthelin, 1880: 35, pl. 26, fig. 28)Pseudonodosaria humilis (Nodosaria humilis Roemer, 1841: 95, pl. 15, fig. 6)Tristix acutangula (Rhabdogonium acutangulum Reuss, 1863: 55, pl. 4, fig. 14)Tristix explanata (T. explanata Epistalié & Sigal, 1963: 62, 63, pl. 29, figs 5, 6)Tristix somaliensis (T. somaliensis Macfadyen, 1935: 11, pl. 1, fig. 7)Tristix temirica (Triplasia temirica Dain, 1934: 29, pl. 3, fig. 25a, b, w)

Morphogroup 5

Geinitzinita kcyniensis (G. kcyniensis Bielecka, 1975: 334, pl. 6, figs 9a, b, 10)Geinitzinita wolinensis (G. wolinensis Bielecka, 1975: 335, pl. 6, figs 11–13)Ichthyolaria nikitini (Frondicularia nikitini Uhlig, 1883: 758, pl. 9, figs 10, 11)Lingulina loryi (Frondicularia loryi Berthelin, 1880: 60, pl. 27, fig. 5)Frondicularia inderica (F. inderica Furssenko & Polenova, 1950: 70, pl. 6, fig. 12a, b)Marginulinopsis bettenstaedti [Lenticulina (Marginulinopsis) bettenstaedti Bartenstein & Brand,1951: 209, pl. 6, figs 144–147]Marginulinopsis buskensis (Marginulina buskensis Bielecka & Po¿aryski, 1954: 56, 57, pl. 8, fig. 39)Marginulinopsis matituna (Cristellaria matituna d’Orbigny, 1849: 242)Marginulinopsis robusta (Marginulina robusta Reuss, 1863: 63, pl. 6, figs 5, 6)Marginulinopsis striatocostata (Marginulina striatocostata Reuss, 1863: 62, pl. 6, fig. 2)Saracenaria alata-angularis (S. alata-angularis Franke, 1936 : 45, pl. 4, figs 13–15)Saracenaria pravoslavlevi (S. pravoslavlevi Furssenko & Polenova, 1950: 45, pl. 4, figs 13–15)Palmula crepidularis (Planularia crepidularis Roemer, 1842: 273, pl. 7B, fig. 4)Palmula malakialinensis [Neoflabellina (Falsopalmula) malakialinensis Epistalié & Sigal, 1963: 56,pl. 26, figs 7–9]Astacolus calliopsis (Marginulina calliopsis Reuss, 1863: 60, pl. 5, fig. 16)Astacolus eritheles (A. eritheles Loeblich & Tappan, 1949: 9, pl. 1, figs 9–13)Astacolus humilis precursoria (Bartenstein & Brand, 1951: 287, pl. 5, figs 126, 127)Astacolus primus franconicus (Cristellaria franconica Gümbel, 1862: 225, pl. 3, fig. 27a, b)

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Astacolus schlönbachi (Cristellaria schlönbachi Reuss, 1863: 65, pl. 6, figs 14, 15)Marginulina declivis (Dentalina declivis Schwager, 1865: 105, pl. 3, fig. 1)Marginulina inaequalis (M. inaequalis Reuss, 1860: 207. pl. 7, fig. 3)Vaginulinopsis embaensis (Cristellaria embaensis Furssenko & Polenova, 1950: 36, pl. 3, figs 9–13)Vaginulinopsis incisiformis (V. incisiformis Bielecka, 1975: 351, 352, pl. 10, figs 7–9)Citharina brevis (Vaginulina brevis Furssenko & Polenova, 1950: 60, pl. 5, figs 3, 4)Citharina paucicostata (Vaginulina paucicostata Reuss, 1863: 65, pl. 4, fig. 8)Citharina raricostata (Vaginulina raricostata Furssenko & Polenova, 1950: 56, pl. 5, figs 5–8)Citharina cf. virgatis (Vaginulina virgatis Furssenko & Polenova, 1950: 62, 63, pl. 4, fig. 1)Citharinella pomeraniae (C. pomeraniae Bielecka, 1975: 330, 331, pl. 6, figs 1, 2)Citharinella portlandensis (C. portlandensis Bielecka, 1975: 331, 332, pl. 6, figs 3–5)Planularia cordiformis (Cristellaria cordiformis Terquem, 1864: 413)Planularia multicostata (P. multicostata Kuznecova, 1960: 29, pl. 2, figs 15–17)Planularia poljenovae (P. poljenovae Kuznecova, 1960: 31, pl. 2, figs 13–15)Planularia subhumilis (Cristellaria humilis Reuss, 1863 65, pl. 6, figs 16, 17)Planularia uilensis (P. uilensis Kuznecova, 1960: 28-29, pl. 2, figs 7, 8)Psilocitharella kochii (Vaginulina kochii Roemer, 1841: 96, pl. 15, fig. 10)Vaginulina contracta (Marginulina contracta Terquem, 1868: 125, pl. 8, figs 13–24)Vaginulina jurassica (Marginulina jurassica Gümbel, 1862: 222, pl. 3, fig. 21a, b)

Morphogroup 6

Lenticulina ambanjabensis (L. ambanjabensis Epistalié et Sigal, 1963: 35, pl. 12, figs 3, 5–6)Lenticulina dogieli (Cristellaria aff. hoplites Wisniowski, T., 1890: Tabl. 8–10)Lenticulina infravolgaensis (Cristellaria infravolgaensis, 1950: Furssenko & Polenova: p. 25, pl. 1,figs 11–14)Lenticulina münsteri (Robulina münsteri Roemer, 1839: 48, pl. 20, fig. 29)Lenticulina nodosa (Robulina nodosa Reuss, 1862: 78, pl. 9, fig. 8)Lenticulina ponderosa (Cristellaria magna Mjatluk, 1939: 52, pl. 3, fig 32a, b, 34a, b)Lenticulina quenstedti (Cristellaria quenstedti Gümbel, 1862: 226, pl. 4, fig. 2a, b)Lenticulina varians (Cristellaria varians Bornemann, 1854: 41, pl. 4 figs 32–34)Lenticulina vistulae elongata (L. vistulae var. elongata Bielecka & Po¿aryski, 1954: 36, pl. 44, fig.16a, b)Lamarckina asteriaformis (L. asteriaformis Kuznecova & Antonova, Antonova et al., 1964: 53, pl. 7,figs 9, 10)Pseudolamarckina polonica [Eponides (Conorbis) polonicus Bielecka & Po¿aryski, 1954: 70, pl. 12,fig. 58 a, b, c]Pseudolamarckina reussi (Lamarckina ? reussi Antonova; Antonova et al., 1964: 50, pl. 8, figs 1–3)

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