GEOLOGICA CARPATHICA, 52, 4, BRATISLAVA, AUGUST 2001
UPPER DEVONIAN—LOWER CARBONIFEROUS FORAMINIFERAL
PALEOBIOGEOGRAPHY AND PERIGONDWANA TERRANES AT THE
Department of Geology and Paleontology, Faculty of Science, Masaryk University, Kotlářská 2, 61137 Brno, Czech Republic;
(Manuscript received December 7, 2000; accepted in revised form June 13, 2001)
Abstract: A new paleobiogeographic subdivision of the Perigondwana terranes is based on the study of Upper Devonian
and Lower Carboniferous calcareous foraminifers. The Brunovistulian, Moesian and Zonguldak terranes in Central and
SE Europe and Asia Minor (the Brunovistulian group of terranes) show close paleobiogeographic relationships to the
East European foraminiferal association. Consequently, they are regarded as a part of the Fennosarmatian Province
which represented a centre for the diversification for most groups of Upper Devonian and Lower Carboniferous calcar-
eous foraminifers. A lower degree of similarity with the East European Platform can be distinguished in the East Avalonian
foraminiferal fauna while that of the Armorican group of terranes displays distinct differences constituting the separate
Armorican Province. The results obtained support the idea that the Brunovistulian group of terranes formed the southern
margin of Laurussia which was colliding with the Armorican group of terranes during the Variscan orogeny. In this
respect, Brunovistulian group of terranes held a geotectonic position similar to that of East Avalonia even though the
accretionary history of the terranes was different.
Key words: Upper Devonian, Lower Carboniferous, Perigondwana terranes, paleobiogeography, calcareous foraminifers.
The most important feature in the evolution of calcareous fora-
minifers is the fact that the highest rates of evolution can be
observed in complicated multilocular forms, while the small
number of morphological features recognized in fossil uniloc-
ular forms makes them inappropriate for such studies.
The first period of differentiation of multilocular, calcareous
foraminiferal faunas can be distinguished in the late Devo-
nian—early Carboniferous. In this period, the benthonic mode
of life in shallow shelf environments predestined calcareous
foraminifers to become a useful tool for paleobiogeographic
studies. To determine the relationships of Perigondwana ter-
ranes, it would evidently be useful to study early Paleozoic
foraminiferal paleobiogeography in detail. However, less di-
versified assemblages of primitive unilocular foraminifers and
the poor knowledge of their distribution makes them inade-
quate for such a study. Nevertheless, even in the Frasnian to
late Tournaisian there seem to be paleobiogeographic differ-
ences between terranes which can have some impact on the
study of relations at the Gondwana-Laurussia interface.
After the break-up of Pannontia many microplates were de-
rived, especially from Gondwana (Erdtmann 2000). The lower
Paleozoic history of Baltica was characterized by the succes-
sive accretion of Perigondwana terranes. However, most of the
paleogeographical models that have been proposed, for exam-
ple, by McKerrow & Scotese (1990), Dalziel (1997), Torsvik
(1996) are too general or concentrated to a certain region. Re-
cent studies of lower Paleozoic paleogeography (e.g. Erdt-
mann 2000; Paris 2000; Robardet et al. 1994; Belka et al.
1997; Ebner et al. 1998; Stampfli 1996; Kozur & Göncüoglu
2000) seem to distinguish a complicated mosaic of terranes at
the Gondwana-Laurussia interface, not only within the Armor-
ican group of terranes, but also in Central Europe and on the
southeastern Laurussian margin. However, most studies are re-
gionally focused and only Erdtmann (2000) included most of
the known terranes in his paleogeographical reconstruction
and summarized the often conflicting state of knowledge.
It is beyond the scope of this paper to discuss the paleobio-
geographical relations of all Perigondwana terranes. Conse-
quently, as the distribution of calcareous foraminiferal fauna in
the upper Devonian and lower Carboniferous was restricted to
shallow shelf environments, only terranes with a microconti-
nental affinity and a rich foraminiferal record will be included.
Terranes of the Caledonian-Variscan orogeny
The most common features which have been applied to dis-
tinguish Perigondwana-derived terranes are the Cadomian
consolidation of the basement, paleomagnetic data, the “Medi-
terranean” faunal affinity or sedimentological criteria (Ordovi-
cian glacigenic sediments). However, all these criteria have
their drawbacks and should be combined (Erdtmann 2000).
Thus, a Cadomian consolidation usually regarded as a part of
the Panafrican consolidation (Ziegler 1986) has recently been
reported in the Urals and Taymir (Puchkov 1998; Glasmacher
et al. 1998). Zelazniewicz (2000) even assumes a Cadomian
orogeny at the southern margin of the Baltica. Even though
such assumptions must still be verified, one has to be more
careful in applying a single criterion for determining a Gond-
There is a growing consensus on the importance of strike-
slip transfer and rotations of terranes which is reflected in the
most recent paleobiogeographic models. This does not only
concern Perigondwana terranes. For example, the Baltica
seems to have been detached from the West-African-South-
American part of Protogondwana, showing a prominent coun-
terclockwise rotation and significant latitudinal transfer during
the early Paleozoic (Erdtmann 2000).
As it was already stressed, in the following contribution, at-
tention will be directed only to discussion of the position of
particular terranes in connection with upper Devonian—lower
Carboniferous foraminiferal paleobiogeography.
In Western Europe, two major groups of terranes between
Baltica (Laurussia) and Gondwana have generally been distin-
1) The East Avalonian terrane group which was accreted to
the southern margin of Baltica during the Caledonian orogeny
and during the Variscan orogeny. It formed the southern prom-
ontory of Laurussia.
2) The Armorican terrane group was accreted to the south-
ern Laurussian margin during the Variscan orogeny. Its major
constituents include the Middle/North Armorican Terrane, the
Central Iberian Terrane, Barrandian and the Saxothuringian
Terrane. Other smaller terranes are also reported (Robardet et
al. 1994; Paris 2000). Some authors also include the Intra-Al-
pine Terrane in this group (Paris 2000).
In Central Europe, a similar situation can be distinguished
(see Fig. 3). The Caledonian accreted terranes include the Lyso-
gory, Malopolska and Brunovistulian terranes (Kalvoda 1995;
Belka et al. 1997). In the Armorican terrane group, the
Saxothuringian and Perunica terranes can be included along
with the Intra-Alpine terranes more to the south and southeast.
Fig. 1. Structural setting of the Brunovistulian, Malopolska and Lysogory terranes. Modified according to Bula et al. (1977). BVT –
Brunovistulian Terrane, MT – Malopolska Terrane, LT – Lysogory Terrane, HKFZ – Hamburg-Krakow Fault Zone, HCF – Holy
Cross Fault, TESZ – Trans-European Suture Zone, MSFZ – Moravo-Silesian Fault Zone, PPFZ – Peri-Pieninian Fault Zone, VDF –
Variscan Deformation Front, CDF – Caledonian Deformation Front, AF – Alpine Front.
UPPER DEVONIAN—LOWER CARBONIFEROUS FORAMINIFERAL PALEOBIOGEOGRAPHY 207
Fig. 2. Tectonic map of the Black Sea region showing the position of the Moesian and Zonguldak terranes. Modified after Okay et al. (1994).
These include the Peri-Mediterranean Terrane and the Noric-
Bosnian Terrane as major units (Neubauer & Handler 2000;
Kováč et al. 1993; Schönlaub & Histon 2000).
In the southeast, at the contact with the East European Plat-
form, the Moesian and Balkan terranes (Yanev 1997) and the
Zonguldak Terrane and other possible terranes in Turkey
(Göncüoglu 1997) can be distinguished. As the Central and
South-Eastern European terranes are not so well known and
have been distinguished only recently, a brief description will
The Brunovistulian Terrane (BT – see Fig. 1) was includ-
ed into the Avalonian group of terranes by Kalvoda (1995).
This terrane has a Cadomian basement and detrital musco-
vites in the lower Cambrian clastics showing the same age
(Belka et al. 2000). Jachowicz & Přichystal (1997) and Fatka
& Vavrdová (1998) have reported Lower Cambrian acritarch
assemblages showing a pronounced similarity with assem-
blages known from the East European Platform. The Cam-
brian trilobite fauna shows affinity to the Baltic Province
(Belka et al. 2000). On the other hand, the middle Devonian
trilobite fauna correlates well with the Barrandian area
(Chlupáč 1969) contrary to the benthonic (coral and brachio-
pod) faunas which are essentially dissimilar to the Barrandian
ones (Galle et al. 1995). It seems that the Brunovistulian Ter-
rane was located close to the Baltica as early as in the Cam-
brian and may has joined the Malopolska Terrane in the early
Devonian (Belka et al. 2000). Important dextral translations
are reported by Grygar (1997).
The Malopolska Terrane (MT – see Fig. 1) is separated
from the Brunovistulian Terrane by a tectonic unit known as
the Krakow-Lubliniec Fault Zone. The Krakow-Lubliniec
Fault Zone is a part of the largely concealed Hamburg-Kra-
kow Fault Zone (see Fig. 1) which is parallel to the Trans-Eu-
ropean Suture Zone (TESZ) and continues southeast of Kra-
kow, where Carpathian flysch is thrust over the Paleozoic
basement (Bula & Jachowicz 1997). According to Belka et al.
(2000), the Malopolska Terrane possesses a Cadomian base-
ment; lower Cambrian sediments yield detrital muscovites
and zircons of both Cadomian and Baltica provenance. Upsec-
tion, the change in provenance from Cadomian to Baltic
sources appears. Belka et al. (2000) thus assumed that the Ma-
lopolska Terrrane was the first Perigondwanan microplate,
which was accreted to the margin of Baltica. Lewandowski
(1992) assumed a significant dextral translation of the terrane
in the Devonian.
The Lysogory Terrane (LT – see Fig. 1) is separated from
the Malopolska Terrane to the south by the Holy Cross Fault.
To the north it is delimited by the TESZ. The basement is un-
known. Middle and Upper Cambrian rocks comprise detrital
muscovites, both Baltic and Cadomian in affinity. It is as-
sumed that the unit had been situated very close to Baltica in
the late Cambrian, even though the Cambrian fauna shows
Gondwanan (Avalonia or Armorica) rather than Baltic affini-
ties (Belka et al. 2000).
In the Moesian Terrane (see Fig. 2), Yanev (1997) reported,
based both on Ordovician acritarchs and trilobites, the pres-
ence of a cool Mediterranean Province. He assumed attach-
ment to the Dobrogea periphery of the Eastern European Plat-
form in the middle to upper Devonian and continuing collision
in the Carboniferous and Variscan collision with the Balkan
Terrane along the Thracian Suture. Crowley et al. (2000) men-
tioned “Cadomian-aged” deformation of arc-related tholeiites
along the northern margins of Central Dobrogea which forms
the northern part of the Moesian Terrane. Ordovician glaciom-
arine sediments in the Balkan Terrane (Yanev 1997a) as well
as Bohemian-type lower Paleozoic fauna (Yanev 1997a) sug-
gest a relationship to the Armorican terranes.
In the Zonguldak Terrane (see Fig. 2) Kozur & Göncüoglu
(2000) reported Cadomian oceanic and island arc sequences
and assumed the lowermost Devonian accretion to the East
European Platform. The presence of Ordovician trilobites of
Welsh affinity may indicate a position close to East Avalonia.
On the basis of the presence of the entirely continental Trias-
sic, they also assume a slightly more northerly position of the
Zonguldak Terrane in comparison with the Moesian Terrane.
This more northerly original position is easily compatible with
the Mesozoic evolution of the Black Sea, a period during
which the Zonguldak Terrane drifted southward (Görür et al.
1997). On the other hand, Sengör (1984) put the Istanbul series
into the Cimmerian continent and Stampfli (1996) regarded
the Zonguldak Terrane (Pontides) as the eastern end of the In-
tra-Alpine Terrane. According to Göncüoglu (1997), both the
Moesian and Zonguldak terranes drifted away from Gondwana
during the Cambrian.
Upper Devonian—Lower Carboniferous
Calcareous foraminifers represent a climatically sensitive
group and their distribution patterns provide useful informa-
tion for reconstructing the paleoclimate and paleogeography.
On this basis, we can distinguish three Devonian-lower Car-
boniferous paleogeographical realms: the tropical/subtropical
Paleotethyan, North American and the boreal Siberian realms
(Mamet & Belford 1968; Lipina 1973; Kalvoda 1990). The
North American realm, even though positioned in the tropical
Fig. 3. Sketch of the microcontinents in the Central European Variscides. Terranes of the Brunovistulian group coloured dark, dotted –
terranes of the Armorican group and light East Avalonia. The relation of enigmatic Lysogory Terrane is not quite clear. The arrows indi-
cate predominating sinistral translation of the East Avalonian Terrane and predominating dextral translation of the remaining Perigond-
wana terranes (see Table 1).
Table 1: Review of Perigondwana terranes at the Laurussia and
Gondwana interface in Central and South-Eastern Europe.
UPPER DEVONIAN—LOWER CARBONIFEROUS FORAMINIFERAL PALEOBIOGEOGRAPHY 209
and subtropical belt, contained a less diversified, but far from
impoverished fauna, in some respects similar to the Siberian
one, probably reflecting the complicated migrational pattern of
foraminiferal taxa (Kalvoda 1990). The relations of the main
realms have been discussed by Kalvoda (1999).
In the original concept of Mamet (Mamet & Belford 1968;
Mamet 1977) the Paleotethyan Realm included both the Lau-
russian and Gondwana shelves. Vdovenko (1980) condition-
ally included the North African margin into the West Europe-
an Province, however, she stated that in the foraminiferal
faunas stenobiont forms with wide paleobiogeographical dis-
tributions predominate and that there are relatively little data.
It is evident that the older paleobiogeographic interpretation
should be revised to incorporate new paleotectonic concepts
distinguishing different terranes with Gondwanan and Baltic
affinities within the Paleotethyan Realm.
In the following, I will try to elucidate the affinity of the
foraminiferal faunas of some Perigondwana terranes. Atten-
tion will be concentrated on those time intervals when paleo-
biogeographical differences within the Paleotethyan Realm
arising from terrane affinity are still apparent (the late Devo-
nian, Tournaisian) and best data are available. Three fora-
miniferal assemblages seem to fit best to this conditions: the
upper Frasnian Multiseptida-Eonodosaria-Eogeinitzina as-
semblage, the upper Famennian Quasiendothyra assemblage
and the upper Tournaisian Kizel-Kosvin assemblage.
Multiseptida-Eonodosaria-Eogeinitzina assemblage (see
We begin our review of the foraminiferal dispersal in the
late Frasnian when the great uniformity of foraminiferal fauna
paralleled the worldwide distribution of reef ecosystems (Kal-
voda 1986) and the first diversification event can be seen. The
Paleotethyan Realm was characterized by the presence of ad-
vanced multi-chambered genera such as Eogeinitzina,
Frondilina, Eonodosaria, Multiseptida, Nanicella and the
primitive Tournayellidae. However, this complete assemblage
is present only on the East European Platform and in the Urals.
Less diversified assemblages can also be found outside the Pa-
leotethyan Realm, both in the North American realm and in
the Omolon-Kolyma and Tian Shan regions (Kalvoda 1990a)
as well as in the Armorican Province.
In the Armorican Province of the Paleotethyan Realm, the
upper Frasnian foraminiferal assemblage has only been de-
scribed in the South Armorican Terrane (Vachard 1994). The
southern Armorican assemblages seem to be far less diversi-
fied than the East European ones. The absence of important
Fig. 4. Simplified paleogeographical map and foraminiferal paleobiogeography during the late Frasnian. L – Lysogory Terrane, M –
Malopolska Terrane, B – Brunovistulian Terrane, M – Moesian Terrane, Z – Zonguldak Terrane, Kaz – Kazakhstan continent, SEU
– Southern Europe, IBR – Iberia, ARM – Armorica, PER – Perunica, EAV – Eastern Avalonia, WAV – Western Avalonia, MEG
– Meguma. Dark lines – boundaries of the faunal assemblages. The shading shows the relation of Perigondwana terranes to the Fen-
nosarmatian Province. Based on the data presented in the text and in the paper by Kalvoda (1990a). North American occurrences of Mul-
tiseptida not included.
genera, Multiseptida and Frondilina as well as primitive
Tournayellidae, is significant.
On the other hand, in the Brunovistulian Terrane, the upper
Frasnian foraminiferal assemblage contains all the multilocu-
lar genera found in the East European Platform and the Urals
(Kalvoda 1990a). This favours the close contact and commu-
nication of this terrane with the East European Platform.
A similar situation is observed in the Moesian Terrane
where upper Frasnian assemblages, very similar to those of
the Eastern European Platform, are reported by Yovcheva
In the Zonguldak Terrane, Dil (1976) reported only Eono-
dosaria from the upper Frasnian assemblage, but only one
outcrop was studied and so the data available may not be rep-
In the Avalonian Terrane, diversified assemblages of the
Upper Frasnian have not been described.
Quasiendothyra assemblage (see Fig. 5)
For the Quasiendothyra assemblage, the successive occur-
rence of Quasiendothyra bella (Chernysheva), Quasiendot-
hyra communis (Rauser) Quasiendothyra regularis Lipina,
and the diversification of Quasiendothyra ex gr. communis
(Rauser), Quasiendothyra kobeitusana (Rauser) and
Quasiendothyra konensis (Lebedeva) are characteristic. The
first diversification of Quasiendothyra and Tournayellidae at
the base of the upper Famennian (the P. marginifera conodont
zone) has been found only in the Paleotethyan Realm. In oth-
er realms the data are not sufficient. The higher upper Fa-
mennian (the P. expansa conodont zone) is marked by the
worldwide dispersal of the Quasiendothyra fauna. This fauna
has also been recorded outside the Paleotethyan Realm, in
the Omolon-Kolyma region, in Alaska as well as in Australia
(Kalvoda 1990a). This migration of Paleotethyan fauna in
both the Southern and Northern Hemispheres suggests cli-
matic warming in this interval (Kalvoda 1986).
In the Armorican Province, Vachard (1988) reported a poor
Quasiendothyra fauna (communication of R. Conil) from the
D-C boundary at La Serre in Montagne Noire while Krylatov
& Mamet (1969) report a richer assemblage in Corsica. The
Quasiendothyra fauna seems to be absent in Spain as well as
in the Pyrenees and in Asturias even though favorauble fa-
cies for such fauna are present (Conil 1977). A more diversi-
fied Quasiendothyra fauna has been described in the eastern
part of the Saxothuringian Terrane by Zukalová (1976).
On the other hand, Conil & Lys (1970) reported rich upper-
most Famennian Quasiendothyra assemblages in the Avalo-
nian Terrane. In the Zonguldak Basin, a Quasiendothyra fau-
na very similar to the eastern European association was
described by Dil (1976). Quasiendothyra assemblages of the
lower and upper part of the upper Famennian, very similar to
the Fennosarmatian occur in the Brunovistulian Terrane
(Kalvoda 1990a). The absence of Quasiendothyra fauna in
the Moesian Terrane is probably due to poor exposure and in
general missing data on upper Famennian foraminifers.
Upper Kizel-Kosvin Assemblage (see Fig. 6)
In the Tethyan Realm, the upper Tournaisian is character-
ized by the diversification and widespread occurrence of the
Upper Kizel fauna (Spinoendothyra, Inflatoendothyra, Eo-
forschia, Dainella, Paraendothyra) which is typical for the
Kizel Cycle of Lipina (1963). It is followed by the succes-
Fig. 5. Simplified paleogeographical map and foraminiferal paleobiogeography during the late Famennian. See explanations in Fig. 4.
UPPER DEVONIAN—LOWER CARBONIFEROUS FORAMINIFERAL PALEOBIOGEOGRAPHY 211
sive occurrence of Dainella, Eotextularia, Tetrataxis, Ur-
banella, Endospiroplectammina, Globoendothyra, Plectogy-
ranopsis, Pseudolituotubella, Eoparastaffellina, Eoendothy-
ranopsis, Eoparastaffella, Omphalotis (Kalvoda 1990a;
Hance et al. 1997) best known from the Kosvin Horizon. The
centre of diversification was again the East European Plat-
form and the Ural Mts. The acme of the migration and prolif-
eration of Paleotethyan calcareous foraminifers seems to co-
incide with the period of the G. typicus-lower S. anchoralis
Zone (Kalvoda 1990). Because the association of several
genera is characteristic of the assemblage, a taxon name was
not applied. The assemblage corresponds broadly to the
Spinoendothyra, Endothyra elegia-Eotextularia diversa and
Eoparastaffella-Eoendothyranopsis zones in Eastern Europe
(see Table 2).
In the Zonguldak Terrane, upper Tournaisian assemblages
showing close relation to the East European Platform have
been described by Dil (1976).
In the Brunovistulian Terrane, upper Tournaisian foramin-
iferal assemblages were reported by Kalvoda (1983, 1990a).
On the basis of a joint study of conodonts, he has shown the
upper Tournaisian age of the fauna regarded, by Conil (1977)
as Visean and he supposes a close relationship to the fora-
miniferal faunas of the East European Platform.
Fig. 6. Simplified paleogeographical map and foraminiferal paleobiogeography during the latest Tournaisian. See explanations in Fig. 4.
Fig. 7. Some important taxa of the described foraminiferal assemblages. 1 – Multiseptida corallina Bykova – Multiseptida-Eonodosar-
ia-Eogeinitzina assemlage; 2 – Quasiendothyra kobeitusana (Rauser) – Quasiendothyra assemblage; 3 – Paraendothyra nalivkini
Tchernysheva – Upper Kizel-Kosvin assemblage; 4 – Eoparastaffella simplex Vdovenko – Upper Kizel-Kosvin assemblage. All
specimens are from the southern part of the Moravian Karst (Czech Republic).
UPPER DEVONIAN—LOWER CARBONIFEROUS FORAMINIFERAL PALEOBIOGEOGRAPHY 213
In the Moesian Terrane, upper Tournaisian foraminifers
have only been described in the Gomotarci borehole. They
correlate well with the Donbas. Therefore the Moesian Plat-
form is included in the Donets-Dobrogea Province (Vdoven-
ko et al. 1981).
The East Avalonian Terrane seems to contain some differ-
ences between the impoverished upper Tournaisian foramin-
iferal fauna of the Dinant Basin and the British SE Province
on one side and the more diversified fauna of the Namur Ba-
sin and especially the British Central Province on the other
(Conil et al. 1979; Kalvoda 1983a). The former regions show
only a weak foraminiferal record in the upper Tournaisian
due to the significant presence of Waulsortian reef facies. In
this respect, we can see some similarities with the southern
Laurentian margin where the Waulsortian Facies is also
widespread (Lane 1982).
A part of the upper Tournaisian foraminiferal fauna of the
Namurian Basin and the British Central Province was original-
ly regarded as “Visean” (Conil & Lys 1964; Conil et al. 1979).
Kalvoda (1983, 1983a) pointed out that the “Visean” assem-
blages of the latter regions as well as of the Campine-Brabant
Basin are partly Tournaisian in age and show a close relation-
ship to the Eastern European faunal associations.
The distribution of foraminiferal faunas in the late Devo-
nian and early Carboniferous of the Paleotethyan Realm
shows important differences between the Armorican terrane
group and the southern Perigondwana terranes (in part the
West European Province of Vdovenko 1980) on one side,
and the East European Platform shelves on the other.
Vdovenko’s (1980) term “West European Province” is mis-
leading for the inclusion of the Avalonian Terrane with its
more diversified fauna. In a similar way, the terms Perigond-
wana or Mediterranean Province could be misleading be-
cause not all Perigondwana terranes could be included within
such a province. Consequently, we prefer to employ the term
Armorican Province to include the Perigondwana terranes
characterized by incomplete foraminiferal phylogeny and
foraminiferal assemblages lower in diversity than the assem-
blages of the East European Platform and the Urals. This re-
gion, which represented a diversification centre of many
groups of late Devonian and early Carboniferous calcareous
foraminifers, is defined here as the Fennosarmatian Province.
The Brunovistulian group of Perigondwana terranes in-
cludes the Brunovistulian, Malopolska, Moesian and Zongul-
Table 2: The correlation chart of the most important late Devonian and early Carboniferous foraminiferal zonations which represents
one of the bases for paleobiogeographical reconstructions. In the chart we can see the correlation of the discussed upper Frasnian, upper
Famennian and upper Tournaisian-lower Visean foraminiferal assemblages. The Paleotethyan Realm includes Namur and Dinant Syncli-
norium, Eastern Europe, Moravia and Tian Shan. Siberian Realm includes Western Siberia, Omolon and Kolyma Massif and Kuznets
Basin. Modified after Kalvoda (1990a).
Fig. 8. Simplified paleogeographical map and groups of Perigondwana terranes based on foraminiferal paleobiogeography. See explana-
tions in Fig. 4.
dak terranes, which show a close similarity to the East Euro-
pean faunal associations and can be regarded as a part of the
Fennosarmatian Province (see Fig. 8). These terranes under-
went a paleogeographical evolution different from the rest of
the Perigondwana terranes – during the Variscan orogeny
they were situated on the southern margins of Laurussia
(Yanev 1997; Göncüoglu 1997; Kalvoda 1995; Belka et al.
1997) in a geotectonic position similar to that of the East
Avalonian Terrane. Thus the presented data support the idea
of continuation of the TESZ SE-wards through the Black Sea
to Asia Minor (Pharaoh et al. 1997). They are not in accor-
dance with the position of the Brunovistulian, Moesian and
Zonguldak terranes in the southern foreland of the Variscan
belt (Matte et al. 1990; Stampfli 1996) and correlation with
the Aquitaine and Montagne Noire Terrane.
The foraminiferal fauna of East Avalonia also shows a
similarity with the East European associations, but some dif-
ferences seem to exist. These are seen in the absence of the
upper Frasnian diversified Multiseptida-Eonodosaria-Eogei-
nitzina assemblage and the presence of Waulsortian reef fa-
cies linking East Avalonia with the southern margin of Lau-
rentia. This may reflect the fact that in contrast to the dextral
translation of the Brunovistulian group and the majority of
Perigondwana terranes, the final accretion of East Avalonia
Acknowledgments: The author is indebted for a thorough
reviewing and useful suggestions to H.G. Herbig, F. Neubau-
er and J. Soták. The study was supported by the research
projects GA205/99/1640 and CEZ-J07/98-143100003.
Arthaud F. & Matte P. 1977: Late Paleozoic strike-slip faulting in
southern Europe and Northern Africa: result of a right lateral
shear zone between the Appalachians and the Urals. Geol. Soc.
Amer. Bull. 88, 1305—1320.
Belka Z., Ahrendt H., Franke W., Bula Z., Jachowicz M. & Wemmer
K. 1997: Accretion of pre-Variscan terranes in the Trans-Euro-
pean Suture Zone: evidence from K/Ar ages of detrital musco-
vites. Terra Nostra 97, 11 21—23.
Belka Z., Valverde-Vaquero P., Ahrendt H., Dörr W., Franke W. &
Wemmer K. 2000: Accretion of first Gondwana-derived ter-
ranes at the marginof Baltica – new data and open questions.
Joint meeting of Europrobe (TESZ) and PACE Projects, Zako-
pane – Holy Cross Mountains, Poland, September 16—23,
Bula Z., Jachowicz M. & Zaba J. 1997: Principal characteristic of
the Upper Silesian Block and Malopolska Block border Zone
(southern Poland). Geol. Mag. 134, 5, 669—677.
Chlupáč I. 1969: Revision of the Middle Devonian trilobites from
Horní Bebešov in the Nízký Jeseník Mts. (Moravia). Sbor.
Geol. Věd, Paleont. 10, 67—103.
Conil R. 1977: The use of Foraminifera for the Biostratigraphy of
the Dinantian in Moravia. In: Wagner R.H. & Holub V.M.
(Eds.): Symposium on Carboniferous Stratigraphy. ÚÚG Pra-
Conil R., Longerstaey P.J. & Ramsbottom W.H.C. 1979: Matériaux
pour l’etude micropaleontologique du Dinantien de Grande
Bretagne. Mém. Inst. Géol. Univ. Louvain 30, 1—187.
Conil R. & Lys M. 1964: Matériaux pour l´etude micropaléon-
tolgique du Dinantien de la Belgique et de la France (Aves-
nois), Algues et Foraminiferes. Mém. Inst. Géol. Univ. Lou-
Conil R. & Lys M. 1970: Données nouvelles sur la Foraminiferes du
Tournaisien inférieur et des couches de passage du Famennien
au Tournaisien dans l´Avesnois. Congres & Coll. Univ. Liege,
55, Coll. Strat. Carbon., 241—265.
Crowley Q.G., Marheine D., Winchester J.A. & Seghedi A. 2000:
Recent geochemical and geochronological studies in Dobro-
gea, Romania. Joint meeting of Europrobe (TESZ) and PACE
Projects, Zakopane – Holy Cross Mountains, Poland, Septem-
ber 16—23, 2000, 1617.
Dalziel I.W.D. 1997: Neoproterozoic-Paleozoic geography and tec-
tonics: Review, hypothesis, environmental speculations. Geol.
Soc. Amer. Bull. 109, 16—42.
Dil N. 1976: Assemblages charactéritiques de foraminiferes du
Dévonien Supérieur et du Dinantien de Turquie (Basin Car-
bonifere de Zonguldak). Ann. Soc. Géol. Belg. 99, 373—400.
Ebner F., Neubauer F. & Rantitsch G. 1998: Alpine and pre-Alpine
terranes in the Alpine-Mediterranean mountain belts: a brief
synthesis. In: Papanikolao D. (Ed.): IGCP project no. 276 Ter-
rane map and terrane descriptions. Ann. Géol. Pays Hellé-
niques 37, 561—573.
Erdtmann B.D. 2000: Neoproterozoic to Ordovician-Silurian Balti-
ca and Laurentia interaction with Protogondwana: critical re-
view of macro and microplate transfer models. Acta Univ.
Carol., Geol. 42, 3—4, 409—418.
Fatka O. & Vavrdová M. 1998: Early Cambrian Acritarcha from
sediments underlying the Devonian in Moravia (Měnín 1 bore-
hole, Czech Republic). Bull. Czech Geol. Survey 73, 1, 65—69.
Galle A., Hladil J. & Isaacson P.E. 1995: Middle Devonian Bioge-
ography of Closing South Laurussia-North Gondwana Varisci-
des: Examples from the Bohemian Massif (Czech Republic),
with emphasis on Horní Benešov. Palaios 10, 221—239.
Glasmacher U., Giesse U., Stroink L., Alekseev A., Reynolds P., Pu-
chkov V. & Walter R. 1998: A Cadomian terrane at the eastern
margin of Baltica – implications for Late Proterozoic paleo-
geography and for the structural evolution of the southwestern
Urals, Russia. 6
Zonenshain Conference on Plate Tectonics
and Europrobe Workshop on Uralides, Moscow, February
1998, Programme and Abstracts, 191.
Göncüoglu M. 1997: Distribution of Lower Paleozoic rocks in the
Alpine terranes of Turkey: Paleogeographic constraint. In:
Göncüoglu M. & Derman A.S. (Eds.): Early Paleozoic Evolu-
tion in NW Gondwana. IGCP Project No 351. II. International
meeting, November 5—11, 1995, Ankara – Turkey. Turkish As-
soc. Petrol. Geol. Spec. Publ. 3, 13—23.
Görür N., Monod O., Okay A.I. Sengör A.M.C., Tüysüz O., Yigitbas
E., Sakinc M. & Akkök R. 1997: Palaeogeographic and tecton-
ic position of the Carboniferous rocks of the western Pontides
(Turkey) in the frame of the Variscan belt. Bull. Soc. Géol.
France 168, 2, 197—205.
Grygar R. 1998: Deformation history of the Variscan accretionary
wedge – Moravosilesian Zone of the Czech Massif. Manu-
script of the habilitation work, VŠB Ostrava (in Czech).
Jachowicz M. & Přichystal A. 1997: Lower Cambrian sediments in
deep boreholes in south Moravia. Bull. Czech Geol. Survey 72,
Hance L., Muchez P., Hou H.F. & Wu X. 1997: Biostratigraphy, sed-
imentology and sequence stratigraphy of the Tournaisian-
Visean transitional strata in South China (Guangxi). Geol. J.
Kalvoda J. 1983: Preliminary foraminiferal zonation of the Upper
Devonian and Lower Carboniferous in Moravia. Knihovnička
ZPN 4, 23—42.
Kalvoda J.1983a: Contribution to the position of the Lower Carbon-
UPPER DEVONIAN—LOWER CARBONIFEROUS FORAMINIFERAL PALEOBIOGEOGRAPHY 215
iferous foraminiferal fauna from Moravia in reconstructions of
the paleobiogeographical dispersal of foraminifers in Europe.
Acta Univ. Carol., Paleont. 4, 329—240.
Kalvoda J. 1986: Upper Frasnian-Lower Tournaisian events and
evolution of calcareous foraminifera, close links to climatic
changes. In: Walliser O.H. (Ed.): Global Bio-events: a critical
approach. Lecture Notes in Earth Sciences. Springer Verlag,
Berlin, 8, 225—236.
Kalvoda J. 1990: Late Devonian—Lower Carboniferous paleobio-
geography of benthic foraminifera and climatic oscillations. In:
Kauffman E.G. & Walliser O.H. (Eds.): Extinction events in
Earth history. Lecture Notes in Earth Sciences. Springer Ver-
lag, Berlin, 30, 183—188.
Kalvoda J. 1990a: Foraminiferal zonation of the Upper Devonian
and Lower Carboniferous in Moravia (Czechoslovkia). Acta
Mus. Moraviae, Sci. Nat. 75, 71—93.
Kalvoda J. 1995: Devonian basins at the eastern margin of Avalonia
in Moravia. Geol. výzk. Mor. Slez. v r. 1994, 48—50.
Kalvoda J. 1999: Tournaisian-Lower Visean calcareous foramin-
ifera: Biostratigraphy and paleogeography. In: Feist R., Talent
J.A. & Daurer A. (Eds.): North Gondwana: Mid Paleozoic ter-
ranes, stratigraphy and biota. Abh. Geol. B.—A. 54, 135—145.
Kováč M., Michalík J., Plašienka D. & Ma o L. 1993: Alpine devel-
opment of the West Carpathians. PřF Masarykovy university,
Brno (in Czech).
Kozur H. & Göncüoglu M. 2000: Main feaures of the pre-Variscan
development in Turkey. Acta Univ. Carol., Geol. 42, 3—4,
Krylatov S. & Mamet B. 1966: Donées nouvelles sur les terrains
paléozoiques de l’Argentella – Tour Margine (Corse). Attri-
bution a la limite dévono—carbonifere du calcaire de Capitello.
Bull. Soc. Géol. France 7, VIII, 73—79.
Lane H.R. 1982: The distribution of the Waulsortian facies in North
America as exemplified in the Sacramento Mountains of New
Mexico. Symposium on the paleoenvironmental setting and
distribution of the Waulsortian facies, El Paso and Alamogor-
do, New Mexico, 96—114.
Lewandowski M. 1992: Paleomagnetic evidence for dextral strike-
slip displacement of the southern block of the Holy Cross Mts
along the East European Platform border during Variscan orog-
eny and its continental-scale geotectonic implications. Geol.
Carpathica, Spec. Issue 43, 151—152.
Lipina O.A. 1963: On the cyclicity of the evolution of Tournaisian
foraminifers. Vopr. mikropaleont. 7, 13—21 (in Russian).
Lipina O.A. 1973: Zonal stratigraphy and paleogeography based on
Tournaisian foraminifers. Vopr. mikropaleont. 16, 3—34 (in
Mamet B.L. 1977: Foraminiferal zonation of the Lower Carbonifer-
ous: methods and stratigraphic implications. In: Kaufmann
E.G. & Hazel J.E. (Eds.): Concepts and Methods of Biostratig-
raphy. Dowden, Hutchinson & Ross, Stroudsburg, 445—462.
Mamet B.L. & Belford D. 1968: Carboniferous Foraminifera,
Bonaparte Gulf Basin, North-Western Australia. Micropaleon-
tology 14, 339—347.
Matte P., Maluski H., Rajlich P. & Franke W. 1990: Terrane bound-
aries in the Bohemian Massif: Results of large-scale Variscan
shearing. Tectonophysics 177, 151—170.
Neubauer F. & Handler R. 2000: Variscan orogeny in the Eastern
Alps and Bohemian Massif: How do these units correlate? In:
Neubauer F. & Höck V. (Eds.): Aspects of Geology in Austria.
Mitt. Österr. Geol. Gesell. 92, 1999, 35—59.
McKerrow W.S. & Scotese C.R. 1990: Paleozoic palaeogeography
and biogeography. Geol. Soc. Mem. 12, 435.
Okay A.I., Sengör A.M.C. & Gorür N. 1994: The Black Sea: kine-
matic history of opening and its effect on the surrounding re-
gions. Geology 22, 267—270.
Paris F. 2000: Early Paleozoic paleobiogeography of northern
Gondwana regions. Acta Univ. Carol., Geol. 42, 3—4, 473—484.
Pharaoh T.C., England R.W., Verniers J. & Zelazniewicz A. 1997:
Introduction: geological and geophysical studies in the Trans-
European Suture Zone. Geol. Mag. 134, 5, 585—900.
Puchkov V. 1998: Cadomides of the Urals and Taymir: connection
with Gondwanan Europe. In: Linnemann U., Heuse T., Fatka
O., Kraft P., Brocke R. & Erdtmann B.D. (Eds.): Prevariscan
Terrane Analysis of Gondwanan Europe. Intern. Geol. Confer-
ence, Excursion Guides and Abstracts. Scr. Staatl. Mus. Miner-
al. Geol. Dresden, 9, 177—178.
Robardet M., Verniers J., Feist R. & Paris F. 1993: Le Paléozoique
anté-varisque de France, contexte paléogéographique et géody-
namique. Géol. France 3, 3—31.
Schönlaub H.P. & Histon K. 2000: The Paleozoic Evolution of the
Southern Alps. Mitt. Österr. Geol. Gesell. 92, 15—34.
Stampfli G.M. 1996: The Intra-Alpine terrane: A Paleotethyan
remnant in the Alpine Varsicides. Eclogae Geol. Helv. 89, 1,
Torsvik T.H., Smethurst M.A., Meert J.G., Voo van der R., McKer-
row W.S., Brasier M.D., Sturt B.A. & Walderhaug H.J. 1996:
Continental break-up and collision in the Neoproterozoic and
Paleozoic – a tale of Baltica and Laurentia. Earth Sci. Rev. 40,
Vachard D. 1988: Calcareous microfossils (Algae, Pseudo-algae and
Foraminifera) from La Serre, Montagne Noire, France. Cour.
Forsch.-Inst. Senckenberg 100, 139—147.
Vachard D. 1994: Foraminiferes et Moravamminides du Givetien et
du Frasnien du Domaine Ligerien (Massif Armorican, France).
Palaeontographica A 231, 1—3, 1—92.
Vdovenko M., Reitlinger A., Yovcheva P. & Spasov C. 1981: Lower
Carboniferous foraminifers of the Gomotarci R-3 borehole
(nortwestern Bulgaria). Paleont., Stratigr. Litol. 15, 3—51 (in
Yanev S. 1997: Paleozoic migration of terrranes from the base-
ment of the eastern part of the Balkan Peninsula from Peri-
Gondwana to Laurussia. In: Göncüoglu M. & Derman A.S.
(Eds.): Early Paleozoic Evolution in NW Gondwana. IGCP
Project No 351. II. International meeting, November 5-11,
1995, Ankara Turkey. Turkish Assoc. Petrol. Geol., Spec.
Publ. 3, 89—100.
Yanev S. 1997a: Paleoclimatic data on terrane movements from
Bulgaria during the Paleozoic. In: Sinha A.K., Sassi F.P. & Pa-
panikolaou D. (Eds.): Geodynamic domains in Alpine-Hima-
layan Tethys. A.A. Balkema, Rotterdam, Brookfield, 347—368.
Yovcheva P.M. 1980: Foraminifers of the Frasnian stage in North-
eastern Bulgaria. Paleont., Stratigr. Litol. 12, 55—68.
Zelazniewicz A. 2000: Rodinian-Baltican link of the Neoproterozo-
ic orogen in southern Poland. Acta Univ. Carol., Geol. 42, 3—
Ziegler P.A. 1986: Geodynamic Model for the Palaeozoic Crustal
Consolidation of Western and Central Europe. Tectonophysics
Zukalová V. 1976: Upper Devonian Stromatoporoids. Foraminifers
and Algae in the borehole Nepasice (Eastern Bohemia). Věst.
Ústř. Úst. Geol. 51, 281—284.