GEOLOGICA CARPATHICA, 49, 2, BRATISLAVA, APRIL 1998
109123
MICROFORAMINIFERS A SPECIFIC FAUNA OF ORGANIC-
WALLED FORAMINIFERA FROM THE CALLOVIAN-OXFORDIAN
LIMESTONES OF THE PIENINY KLIPPEN BELT
(WESTERN CARPATHIANS)
MILAN MIÍK
1
and JÁN SOTÁK
2
1
Department of Geology and Paleontology, Faculty of Sciences, Comenius University, Mlynská dolina, 842 15 Bratislava, Slovak Republic
2
Geological Institute, Slovak Academy of Sciences Bratislava; Branch: Severná 5, 974 01 Banská Bystrica, Slovak Republic
(Manuscript received May 11, 1997; accepted in revised form March 12, 1998)
Abstract: The organic linings of minute foraminifers can be naturally stained by Fe-oxides during early diagenesis.
This protects them against destruction and make them visible in thin sections. Up till now the microforaminiferal
linings were described only from extractions for palynological investigation. A set of morphotypes of the foramin-
iferal linings from thin sections of Callovian-Oxfordian limestones and Lower Cretaceous cherts is illustrated with
comments. The morphological aspects of the foraminiferal linings allow us to associate them into morphogroups and
form genera. Some of well-developed linings are sufficient for their generic as well as subgeneric classification
enclosing more than 30 foraminiferal taxa. The reasons for the organic-walled structure of microforaminifers are
inferred in environmental stress or dissolution of rigid tests after death. The possibility of using some linings in
foraminiferal stratigraphy is discussed.
Key words: Western Carpathians, Norian, Callovian-Oxfordian, Hauterivian-Barremian, organic linings, foraminifers.
State of preservation
Microforaminiferal linings are rarely preserved; in such
favourable cases they can be extracted by the palynological
preparation procedure. Some linings were also isolated by
the cautious dissolution of the recent foraminifers by very
diluted acids (Taugourdeau-Lantz & Poignant 1964; Loebli-
ch & Tappan 1964; Cohen & Guber 1968). Up till now they
were almost always reported from palynological prepara-
tions, from the insoluble residues, therefore we want to
present here the first larger collection from thin sections of
limestones.
The foraminiferal linings isolated by the palynological pro-
cedure can be well stained by safranin (Stancliffe 1989). Our
findings are based on the fact that under the favourable condi-
tions they can be naturally stained to a red colour by diage-
Introduction
The term microforaminifers was introduced by Wetzel
(1957) for linings of juvenile parts of foraminiferal tests
chitinous membranes. The first mention concerning these
objects is in the paper of Wilson & Hofmeister (1952). They
found them in the palynological preparations. Due to their
minute dimensions (according to most authors below 150
µ
m) the term microforaminifers became usual, but many au-
thors considered it unsuitable and other synonyms were pro-
posed: Scytinascia (Deák l964; Courtinat 1979), basales
membranes (e.g. Taugourdeau-Lantz & Poignant 1964),
microforaminiferal linings (e.g. Stancliffe 1989), chitinous
linings (Tappan & Loeblich 1965), nannoforaminifers, pa-
lynoforaminifers (Pantiæ & Bajraktareviæ 1988), tapeta of
foraminifers (e.g. Pacltová 1978).
Fig. 1. Location of the cited localities with microforaminifers in the Western Carpathians.
110 MIÍK and SOTÁK
netic impregnations of Fe-oxides and become visible in thin
sections of limestones. The impregnation took place in an
early stage of diagenesis and is a conservation agent condi-
tioning the preservation of the microforaminiferal linings. A
huge majority of non-impregnated microforaminiferal mem-
branes are obviously destroyed during diagenesis. They are
however abundant in thin sections from the studied red or
reddish limestones (Pl. I: Fig. A), although part of the calcitic
tests was dissolved. Their organic membranes attracted and
concentrated the migrating Fe-oxides and acquire a deep red
colour in contrast to the neighbouring parts of the limestone.
Occurring rarely in the grey limestones they are also stained
red, because they can trap the Fe-oxides from the very diluted
solutions. We did not succeed in extracting the iron-impreg-
nated linings using the conventional palynological methods.
They probably lose their elasticity as a result of the impregna-
tion and break down easily during the preparation.
Several microforaminiferal linings were found in thin sec-
tions from chert nodules in the Hauterivian-Barremian
Muráò Limestone (Pl. VII: Figs. K, L) and chert nodule in a
Tithonian pelagic limestone (Pl. VII: Fig. M); in those cases
they were not impregnated by Fe-oxides. Silicites are
known to be favourable for the preservation of microfossils
consisting of organic compounds; e.g. in thin sections of the
mentioned locality Muráò. Besides microforaminiferal lin-
ings pollen grains and Fungisporonites occurred there (Mi-
ík 1990, Pl. III, IV).
The disadvantage of thin section studies of microforamin-
iferal linings (non-oriented sections) is counterbalanced by
some advantages. Unlike palynological preparation (centrif-
ugal procedure) specimens are not broken or additionally
deformed, larger specimens are not eliminated by the sepa-
ration of fine fraction (several palynologists use fraction be-
low 0.0025 mm), some sections can render an indication of
the internal structure of the test.
Possible reasons for the tiny dimensions
of microforaminifers
The very small dimensions of the chitinous membranes
are peculiar. The most common explanation is that they rep-
resent only the juvenile part of the foraminiferal test. How-
ever they could represent dwarf forms of adult individuals
living under unfavourable conditions. It should be stressed
that a quarter of the specimens illustrated here are from nep-
tunian dykes filled by red micrite, others occurred in the
condensed facies of ammonitico-rosso type. It is surpris-
ing that in our thin sections with abundant linings of juve-
nile tests almost no large adult representatives of the same
species were found. Courtinat (1989) also stressed the fact
that the found microforaminiferal types correspond neither
to the foraminiferal taxa cited from the Oxfordian of the
Jura Mts., nor to families known for the presence of organic
linings in their tests. The lack of adult individuals corre-
sponding to the embryonic stages could also be explained
by the adult specimens living in different environments or
their calcitic tests being entirely dissolved during early di-
agenesis, as suggested by the dissolution of younger cham-
bers without organic linings (Pl. II: Figs. AL), this seeming
to be the most plausible explanation. The abundance of mi-
croforaminiferal linings and the rarity of adult specimens
might reflect the well-known fact that in some animal
groups only a small percentage of juveniles survive to the
adult age. There is another theoretical possibility that some
species could remain during all their lives in the juvenile
state like the well-known Mexican Amphibian Axolotl. A
somewhat similar solution was proposed by Deák (1964);
according to her Scytinascia could have possessed an exclu-
sively organic test. However we observed numerous cases
with the red stained membrane only in the juvenile part of the
preserved calcitic foraminiferal test (Pl. II: Figs. AL).
Hence, such a generalization should be discarded. The or-
ganic lining gradually thins towards the younger chambers
as was noted by previous authors, too (Stancliffe 1989,
p. 350; Pantiæ & Bajraktareviæ 1988, p. 956). In exception
cases the lining may also be present in the adult forms of
some agglutinated foraminifers (e.g. Spiroplectinella, Zlin-
ská & Ètyroká 1993).
Stratigraphic horizons with microforaminifers
Microforaminiferal linings were found in the sediments
from the Permian to the Quaternary. Stancliffe (1989, text-
figure 5) summarized the papers of more than 60 authors in
a table. Most of the cited authors described them from the
Upper Jurassic which is also the age of our specimens. We
can complete Stancliffes list by some more authors who
cited and illustrated foraminiferal linings: Soták (1986, p.
3031, Pl. 5) from neptunian dykes of Upper Triassic? lime-
stones (blocks in conglomeratic flysch of the Magura Unit,
Plate I: Microforaminiferal linings (basales membranes) im-
pregnated by Fe-oxides. Fig. A Abundant occurrence in the
red nodular Czorsztyn Limestone with filament microfacies (ju-
venile planktonic bivalves Bositra). Oxfordian, Czorsztyn Succes-
sion, Pieniny Klippen Belt, Kyjov-Pusté Pole. Thin section No.
211127,
×
40. Fig. B Microforaminifer in the channel of a
crinoidal columnalium. Upper Tithonian filling of a neptunian
dyke. Kyjov-Pusté Pole. Thin section No. 19440,
×
80. Fig. C
Morphotype of buliminid foraminifer (aff. Dentalinopsis oolithica
(Terq.)). Red micrite from a neptunian dyke synchronous with the
surrounding Callovian limestones, klippe Babina near Bohunice.
Thin section No. 12385,
×
80. Fig. D Morphotype of verneulinid
foraminifer (aff. Verneuilinoides minuta Said & Barakat). At the
difference to the preceding cases it was not naturally stained. Nod-
ular chert from the Muráò Limestone, Upper Hauterivian-Lower
Barremian, southern wall of Muráò peak, Belanské Tatry Mts.
Thin section No. 12520,
×
120. Fig. E Morphotype of trocham-
minid foraminifer (aff. Trochammina squamatoformis Kaptarenko-
Chernousova). Organic membrane is finelly perforated. Callovian-
Oxfordian limestones of the Vratec-castle klippe. Thin section
No. 11494,
×
150. Fig. F Morphotype of trochamminid foramin-
ifer with granular structure of organic membrane. Callovian lime-
stone, klippe Kostelec near Povaská Bystrica, Kostelec Succes-
sion. Thin section No. 21205,
×
100. Fig. G Morphotype of
textularid foraminifer (aff. Bigenerina jurassica (Haeusler)). Call-
ovian-Oxfordian filling of a neptunian dyke, Vratec-castle klippe.
Thin section No. 21095,
×
100.
→
PLATE I 111
112 PLATE II
MICROFORAMINIFERS A SPECIFIC FAUNA OF ORGANIC-WALLED FORAMINIFERA 113
Western Carpathians), Derman et al. (1995, Pl. 1, Fig. 19)
from the Middle Jurassic of Turkey; Èorná (1972, Pl. 11,
Figs. 49) from the Albian of the Manín Unit, Western Car-
pathians; Èorná in Grün et al. (1972, Pl. 22, Figs. 810)
from the Aptian of the Flysch Belt, Eastern Alps, Wiener-
wald; Snopková & Samuel (1981, Pl. LIII, Fig. 3) from the
Upper Senonian to Paleocene of the Dukla Unit, Flysch
Belt, Western Carpathians; Pacltová (1978, Pl. 1, Fig. 3) and
ítt et al. (1997, Pl. 5, Figs. 13, 9) from the Cretaceous of
the Bohemian Massif; Holcová et al. (1996, Tab. 5, Fig. 12)
from the Lower Badenian, South Slovakia; Head (1993, Fig.
5, 14) from the Pliocene of England.
We present in this paper with a kind permission V. Sitár his
non-published material extracted for palynological studies
from the Norian Hauptdolomite in the basement of the Slovak
part of the Vienna Basin (Pl. VII: Figs. AG) and from the
Posidonia marls (Podzamcze Formation), Middle Jurrassic,
Slovak part of the Pieniny Klippen Belt (Pl. VII: Figs. HJ).
Descriptive nomenclature of microforaminifers
Deák (1964) proposed the term Scytinascia for the whole
group of microforaminiferal linings and also formal names
for some genera and species (e.g. Ormathascia covering
straight uniserial tests of Nodobacularia type; Trochiliascia
for planispiral or trochospiral tests of Globigerinoides type;
Rhodonascia for the tests of Trochammina type etc.). Tappan
& Loeblich (1965) rejected this procedure and tried to at-
tribute some of these objects to existing families and with a
question-mark up to the generic level. The term Scytinascia
was rarely used (e.g. Courtinat 1989; Courtinat & Méon
1991); nobody is now introducing of new names for similar
artificial taxa. Courtinat (1989) and Stancliffe (1989) used in-
formal classification based on the morphotypes. We do not
consider it useful to apply the terms of Pantiæ & Bajraktareviæ
(1988), either palynoforaminifers for the foraminifers
found in the macerate for palynomorph studies, or nannofor-
aminifers s. str. for tiny foraminifers in the preparations for
the study of calcareous or siliceous nannoplankton. In such a
way it would be necessary to designate our material as thin-
section microforaminifers.
Comments on microforaminifers presented
from the Western Carpathians
The large majority of the specimens reproduced on the
plates comes from red Callovian-Oxfordian limestones,
mainly of the Czorsztyn Succession of the Pieniny Klippen
Belt, rarely from the Manín Succession (Pl. I: Fig. D; Pl. VI:
Figs. H,J; Pl. VII: Figs. K,L), Kostelec Succession (Pl. I:
Fig. F; Pl. III: Fig. J), from the Liassic of the Niná Succes-
sion (Pl. II: Fig. G) and from pebbles of rocks in the Pale-
ocene Proè Conglomerate (Pl. III: Figs. C,F; Pl. IV: Figs.
E,F; Pl. V: Fig. G; Pl. VI: Fig. L). For the situation of locali-
ties see Fig. 1.
As an example of the associations with foraminiferal lin-
ings from the Callovian-Oxfordian limestones a thin section
description is given (locality Kyjov-VIIa): Red biomicrite-
packstone with filaments foraminifer microfacies. The
main constituent is filaments thin juvenile bivalve shells
of Bositra sp., non-oriented (bioturbation), rarely also frag-
ments of thicker shells perforated by boring algae. Abundant
foraminifers belong to the genera Trocholina, Patellina, Oph-
thalmidium, Globuligerina, and to the foraminiferal linings
(Pl. I: Fig. A). Less frequent are echinoderm ossicles, ostra-
codes, sponge spicules, mainly rhaxa, filled by calcite aggre-
gate, gastropods and exceptionally holothurian sclerites
Theelia sp. A slight admixture of clastic quartz (silt size); mi-
crostylolites stained in red and small irregular concentrations
of Fe-oxides are found.
The suspicion that sometimes not only the chitinous linings
but also the calcitic wall of foraminiferal tests was replaced by
iron oxides can be discarded by their simultaneous presence
in the same thin sections visible on the Pl. II. Several unstained
microforaminiferal linings are also figured here for the purpose
of comparison. They come from thin sections of black nodular
cherts in Barremian Muráò Limestone (Pl. VII: Figs. K,L) and
Plate II: Microforaminiferal linings (basale membranes) from
the Callovian-Oxfordian limestones, Czorsztyn Succession, Pi-
eniny Klippen Belt. Figs. AG Organic membranes developed
only in the juvenile parts of the foraminiferal tests. They were nat-
urally stained by Fe-oxides (black on the photos). Last chambers
were dissolved; several of them are still visible as ghosts. All
specimens are from the Callovian-Oxfordian limestones of the
Czorsztyn Succession, Pieniny Klippen Belt (except Fig. G).
Figs. AD Morphotype of involutinid foraminifers (Trocholina
gracilis Blau). Limestone in the neptunian dyke filling. Klippe Ba-
bina near Bohunice. Fig. A Thin section No. 12497,
×
150. Fig.
B Klippe Kyjov-Pusté Pole. Thin section No. 21085,
×
100. Fig.
C Klippe Krasin near Dolná Súèa. Thin section No. 21527,
×
50.
Fig. D Klippe Kyjov-Pusté Pole. Thin section No. 21101,
×
100.
Figs. EF Morphotype of ammodiscid foraminifers (Glomo-
spirella otorica Romanova). Fig. E Klippe in the Boleov Val-
ley. Thin section No. 19347,
×
160. Fig. F Neptunian dyke fill-
ing, Vratec-castle klippe. Thin section No. 8536,
×
60. Fig. G
Morphotype of spirillinid foraminifer (Semiinvoluta violae Blau).
Middle Liassic limestone, Niná Succession, klippe Lutý Potok.
Thin section No. 21177,
×
100. Fig. H Uncertain foraminiferal
linings. Neptunian dyke filling, Vratec-castle klippe. Thin section
No. 21091,
×
100. Fig. I Morphotype of trochamminid foramini-
fer (aff. Trochammina pulchra Ziegler), the same locality. Thin
section No. 21096,
×
100. Fig. J Morphotype of textularid fora-
minifer (aff. Textularia ripleyensis Berry). As above. Thin section
No. 21080,
×
60. Fig. K Morphotype of verneulinid foraminifer
(Verneuilinoides minuta Said & Barakad), neptunian dyke filling,
klippe Kyjov-Pusté Pole. Thin section No. 21085,
×
120. Fig. L
Morphotype of trochamminid foraminifer, last chamber as a ghost,
the same locality. Thin section No. 21107,
×
100. Figs. MS
Morphotype of dentaliferous species of Reophacidae Hor-
mosinidae, all samples from the Callovian-Oxfordian limestones,
Czorsztyn Succession. Fig. M Scherochella aff. minuta (Tap-
pan), Vratec-castle klippe. Thin section No. 5875,
×
140. Fig. N
Nodulina aff. dentaliformis (Brady), thin section No. 5058,
×
140. Fig. O Reophax aff. chrysalis (Haeusler), klippe Babina
near Bohunice. Thin section No. 12479,
×
230, Figs. PR
Reophax aff. scorpiurus Montford, Fig. P Klippe Kyjov-Pusté
Pole. Thin section No. 23991,
×
200. Fig. R Klippe Drieòová
near Krivoklát. Thin section No. 10792,
×
185. Fig. S Pseudon-
odosinella aff. nodulosa (Brady), compressed specimens, the same
locality. Thin section No. 10622,
×
185.
←
114 MIÍK and SOTÁK
specimens extracted by palynological procedure from the No-
rian Hauptdolomite with shaly intercalations (Pl. VII: Figs. A
G) and Bajocian Supraposidonia shales (Podzamcze Forma-
tion Pl. VII: Figs. HJ); both kindly provided by V. Sitár.
The basal membranes (linings) from foraminiferal tests dis-
solved during early diagenesis are rarely wrinkled by drying
and compaction (Pl. I: Figs. C,D). Sometimes they possess a
granular (pitted) structure reflecting the uneven internal sur-
face of tests agglutinated of larger particles (Pl. I: Figs. E,F).
The necks between the chambers in uniserial and biserial
types (Pl. I: Fig. G; Pl. II: Fig. N) are rarely preserved.
The size of all studied microforaminifers varies between
68
µm and 144 µm.
Typology of foraminiferal linings
and their systematic classification
The morphologies of foraminiferal linings allow us to as-
sociate them into morphogroups and form genera, some
forms could also be attributed to known taxa of Jurassic or
Early Cretaceous species (cf. Haeusler 1890; Mjatljuk
1939; Wicher 1938; Dain 1972; Bartenstein & Brand 1937,
1951; Loeblich & Tappan 1950; Bielecka & Pozaryski
1954; Kaptarenko-Chernousova 1959; Seibold 1960; Neagu
1972; Kuznetsova & Gorbachik 1985; Blau 1987a,b; Weidlich
1990; Nagy & Johansen 1991; Neagu & Neagu 1995, etc.).
Morphotypes of textularid, bolivinid and buliminid foraminifers
The linings of this morphogroup are triangular shape, biser-
ially coiled and globularly or cuneately chambered. Series of
chambers are joined by narrow necks forming a zigzag prolif-
eration between them (e.g. Pl. VII: Figs. C,D). Such morpho-
types of lininigs respond to the Stancliffes oblique necks
giving a zigzag appearence in his biserial type I. Some of
them are also consistent with Courtinats (1989) biserial type
S2. Among linings of this morphogroup the four main form
genera are recognizable Textularia (aff. T. anglica Lalicke
Pl. IV: Figs. IS; T. ripleyensis Berry Pl. II: Fig. J), Bo-
livina (aff. B. rhumbleri Franke Pl. IV, Figs. AC),
Pseudobolivina (aff. P. variana Eicher Pl. IV: Figs. EH;
Pl. VII: Figs. CE; P. clavellata (Loeblich & Tappan)
Pl. III: Fig. B) and Dentalinopsis (aff. D. oolithica (Terq.)
Pl. I: Fig. C). This morphogroup also includes biserial forms
later abruptly reduced to a uniserial and rectilinear stage with
terminal aperture. Considering the biseriallity of these forms
they could not be triserial ataxophragmiids (e.g. Gaudryinel-
la), but probably a textularid species of the genus Bigenerina
(B. jurassica (Haeusler) Pl. I: G; Pl. III: I,J a species
with wholly uniserial and not lax-uniserial adult stage as in
the emended species of Bicazammina Neagu & Neagu 1995)
and the genus Haghimashella (H. arcuata (Haeusler), emend.
Neagu & Neagu 1995 Pl. III: Figs. GH; Pl. V: Fig. O).
Morphotypes of ataxophragmiid and verneulinid foraminifers
The linings of this morphogroup occur as trochoidal tests
with triserial chambers and interiomarginal aperture. They are
developed as triserial high-spired forms (Verneuilinoides
minuta Said & Barakat Pl. I: Fig. D; Pl. II: Fig. K), conical
forms coiled in 23 whorls (Duotaxis metula Kristan Pl.
V: Fig. 6, Jurassic foraminifers originally referred as Tet-
rataxis are thought to be a synonymum of Duotaxis
Fugagnoli 1996) and forms with sutures depressed in growth
axis (Belorussiella aff. bolivinaeformis Akimets Pl. III:
Figs. EF).
Morphotypes of trochamminid, haplophragmoid and lituol-
id foraminifers
This morphogroup encloses a number of lininigs registrat-
ed in the thin sections as well as in the palynological prepara-
tions. Some of the low-spired linings in our thin sections
show a close similarity to the species of Trochammina squa-
matoformis Kaptarenko-Chernousova (Pl. I: Fig. E), Tro-
chammina pulchra Ziegler (Pl. II: Fig. I) and Trochammina
globoconica Tyszka & Kaminski (Pl. V: Fig. D). Small Glo-
bigerina-shaped linings (Pl. V: Figs. AB) of this morpho-
group correspond to the trochamminid species Ammoglobige-
rina canningensis (Tappan) and A. globigeriniformis (Parker
& Jones). Other specimens with subcircular outline, planispi-
ral whorls and trapezoid-shaped chambers could be attributed
to the form genera Recurvoides (Pl. VI: Figs. A, CD, EG,
K). Among the thin section microforaminifers there are also
linings resembling in their appearance Ammobaculites
(A. irregularis (Gümbel)), Bulbobaculites (B. oviloculus
Nagy & Johansen) and Haplophragmium sp.
Plate III: Microforaminiferal linings (basale membranes) from
the Callovian-Oxfordian limestones, Czorsztyn Succession, Pi-
eniny Klippen Belt (except Figs. C,F). Fig. A Lining of den-
taliferous species of Hormosinidae (aff. Scherochella minuta
(Tappan)). Klippe Krasin near Dolná Súèa. Thin section No.
21117,
×
80. Figs. BC Morphotype of bolivinid foraminifers.
Fig. B Pseudobolivina aff. clavellata (Loeblich & Tappan).
Thin section No. 19558,
×
120. Fig. C Pseudobolivina aff. vari-
ana Eicher, pebble of the Liassic limestone in the Paleocene Proè
Conglomerate, Klippen Belt. Beòatina-II-108. Thin section No.
17167,
×
185. Fig. D dentaliferous lining of Reophacidae
(aff. Reophax scorpiurus Montfort), klippe Krasin near Bohunice.
Thin section No. 19487,
×
100. Figs. EF Morphotype of ver-
neulinid foraminifers with sutures depressed in growth axis (aff.
Belorussiella bolivinaeformis Akimets) Fig. E Klippe Babina
near Bohunice. Thin section No. 20181,
×
160. Fig. F Thin sec-
tion No. 17168,
×
185. Figs. GH Linings of textularid species
Haghimashella arcuata (Haeusler), Vratec-castle klippe. Thin
section No. 7005,
×
136. Fig. H Klippe Kyjov-Pusté Pole. Thin
section No. 17187,
×
100. Fig. IJ Linings of textularid species
Bigenerina jurassica (Haeusler). Fig. I Neptunian dyke filling,
Vratec-castle klippe. Thin section No. l222l,
×
l50. Fig. J
Klippe Kostelec, Kostelec Succession. Thin section No. ll633,
×
185. Figs. KM Morphotype of nubeculariid foraminifers
(Nubecullinella infraoolithica (Terq.)) Fig. K Klippe Krasin near
Dolná Súèa. Thin section No. 23892,
×
200. Fig. L Klippe Kyjov-
Pusté Pole. Thin section No. 23991,
×
170. Figs. MN Morpho-
type of nubeculariid? foraminifers. Fig. M Klippe Monèeková
near Èervený Kameò. Thin section No. 10797,
×
270. Fig. N
Klippe Babina near Bohunice. Thin section No. 20191,
×
160.
→
PLATE III 115
116 PLATE IV
MICROFORAMINIFERS A SPECIFIC FAUNA OF ORGANIC-WALLED FORAMINIFERA 117
The foraminiferal linings extracted from the Upper Triassic
(Pl. VII: Figs. A,B) and Middle Jurassic sediments (Pl. VII:
Figs. H,J) and in the thin sections of silicites (Pl. VII: Figs.
KM) show the same appearance of linings illustrated from
the palynological preparations. In Stancliffes nomenclature
they are linings of planispiral type III and IV. Courtinat
(1989) quoted them from the Oxfordian as PS type. Derman
et al. (1995, Pl. I: Fig. 19) illustrated them from the Middle
Jurassic. Deák (1964, Pl. X, Figs. 1719) cited them from the
Albian under the name Trochiliascia cuvillieri; Macko
(1963) from the Turonian under the name Knasteria spiralis,
Èorná (1972, Pl. XI: Figs. 4,8) from the Albian, Èorná in
Grün et al. (1972, Pl. 22: Figs. 810) from the Aptian; Sidó
(1975) equally from the Aptian as species of Globigerinel-
loides algerianus Cushman & Ten Dam and Hedbergella sp.,
Pacltová (1978, Pl. I: Fig. 3) from the Upper Cretaceous;
Pantiæ & Bajraktareviæ (1988, Pl. I: Figs. 1123; Pl. II: Figs.
410) from the Cenomanian; Head (1993, Fig. V: 14) from
the Pliocene.
Morphotypes of our extracted linings (Pl. VII: Figs. AB,
EM) show an ovate proloculus followed by planispirally or
low-trochospirally enrolled, evolute or slightly embracing
coils. The chambers are piriferous, lobate on periphery,
joined by intracameral necks on the umbilical side. Sutures
are deeply depressed, backwards and expanding from the
outer wall. The features described are diagnostic for a wider
range of foraminiferal taxa. Nevertheless, they should be
rather agglutinated foraminifera which normally secreted
thick organic inner layers with a high iron, sulphur and mag-
nesium content (e.g. Trochamminacea Brönnimann &
Whittaker 1988). Considering the morphological aspects
and agglutinated nature of these linings, they could be at-
tributed to the form genera Haplophragmoides (aff. H.
concavus (Chapman) Pl. VII: Figs. AB, KM; aff. H. glo-
bigerinoides (Haeusler) Pl. VII: Figs. FG) and Trocham-
mina (aff. T. inflata (Montagu) Pl. VII: Figs. HJ).
Morphotypes of involutinid, ammodiscid
and spirillinid foraminifers
This morphogroup encloses conical or discoidal forms with
trochospiral, planispiral and/or streptospiral coiling. Based on
coiling type, there are the wholly trochospiral forms of the
genus Trocholina (T. gracilis Blau Pl. II: AD), wholly
planispiral forms of the genus Spirillina (S. elongata Bielec-
ka & Pozaryski Pl. V: Fig. H) and Ammodiscus (A. vari-
ans Kaptarenko-Chernousova Pl. V: Fig. G), faintly tro-
chospiral forms with shallow umbilical side of the genus
Semiinvoluta (S. violae Blau Pl. II: F), early streptospiral
and later planispiral forms of the genus Glomospirella (G.
otorica Romanova Pl. II: Figs. EF; G. reata (Eicher)
Pl. V: Fig. P) and trochospiral forms close to the genus
Arenoturrispirillina (Pl. V: Fig. H). The organic linings of
these forms are commonly reduced on juvenile parts of the
calcareous tests (e.g. Pl. II: Figs. B,D,E).
Morphotypes of dentaliferous foraminifers
An uncertain morphogroup of rectilinear forms with uniser-
ial chambers almost non-overlapping or slightly overlapping,
mostly spheric or strongly elongated (Pl. II: Figs. MS; Pl.
III: Fig. A). The maximal number of eight chambers displays
the specimen Pl. III: Fig. A with a small proloculus. Due to
the lack of outer tests the attribution of these linings is not so
evident. Some of them show appearance of Nodosariidae.
However, the co-occurrence of dentaliferous linings with
true nodosariids makes it possible to attribute them to another
taxa, most probably to multilocular forms of Reophacidae
and Hormosinidae (Reophax aff. helveticus (Haeusler) Pl.
III: Fig. D; R. aff. scorpiurus Montfort Pl. II: Figs. PR;
R. aff. chrysalis (Haeusler) Pl. II: Fig. O; Nodulina aff.
dentaliformis (Brady) Pl. II: Fig. N; Scherochella aff.
minuta (Tappan) Pl. III: Fig. A; Pl. II: Fig. M; Pseudono-
dosinella aff. nodulosa (Brady) Pl. II: Fig. S), which as
agglutinated foraminifera secreted exclusively chitinous lin-
ings (Jakovleva 1979). This attribution is also supported by
the raised apertural necks between successive chambers (Pl.
III: Fig. D). Similarly, some other foraminiferal linings are
also comparable to Reophacidae. Such linings are shown on
Pl. IV: Figs. MN having a large sphaerical proloculus (mac-
rosphaerical forms Fig. M) or plani/streptospiral segment
(microsphaerical forms Fig. N) followed by helicoidal
stage with several loops (max. 6) around the axis of the col-
umella. These meandrospiroid? and later tubiform specimens
(Pl. IV: Figs. MN) recall the Triassic representatives of Tur-
riglominidae Zaninetti (from the Lower Cretaceous they
Plate IV: Microforaminiferal linings from the Callovian- Oxfor-
dian limestones, Czorsztyn Succession, Pieniny Klippen Belt (ex-
cept Figs. E,F,J). Figs. AC Morphotype of bolivinid foramini-
fers (aff. Bolivina rhumbleri Franke). Fig. A Klippe Sivá Skala
near Èervený Kameò. Thin section No. 21183,
×
60. Fig. B Nep-
tunian dyke filling, Vratec-castle klippe. Thin section No. 21080,
×
120. Fig. C Klippe Kyjov-Pusté Pole. Thin section No. 19438,
×
240. Figs. DH, JL, OR, TU Morphotype of bolivinid fora-
minifers (aff. Pseudobolivina variana Eichler). Fig. D Klippe
Kyjov-Pusté Pole. Thin section No. 18018,
×
80. Fig. E Liassic
limestone pebble from the Paleocene Proè Conglomerate, Beòatina-
II-108. Thin section No. 17168,
×
185. Fig. F As above. Thin sec-
tion No. 16443,
×
180. Fig. G Klippe Krasin near Dolná Súèa.
Thin section No. 21098,
×
160. Fig. H Klippe Kyjov-Pusté Pole.
Thin section No. 21107,
×
120. Fig. J Klippe Kostelec, Kostelec
Succession. Thin section No. 11633,
×
185. Fig. K Klippe Babina
near Bohunice. Thin section No. 12479,
×
115. Fig. L Klippe
Drieòová near Krivoklát. Thin section No. 9614,
×
130. section No.
10792,
×
185. Fig. O As above. Thin section No. 11379,
×
185.
Fig. P Neptunian dyke filling, klippe Babina near Bohunice. Thin
section No. 12479,
×
100. Fig. R Klippe Kyjov-Pusté Pole. Thin
section No. 2399l,
×
200. Fig. T Neptunian dyke filling, Vratec-
castle klippe. Thin section No. 23383,
×
145. Fig. U Neptunian
dyke filling, klippe Babina near Bohunice. Thin section No. 12479,
×
230. Figs. I,S Morphotype of textularid foraminifers (aff. Textu-
laria anglica Lalicker). Fig. I Klippe near Krivoklát. Thin sec-
tion No. 12221,
×
150. Fig. S Klippe Hrebeò near Èervený Ka-
meò. Thin section No. 2393l,
×
200. Figs. MN Morphotype of
Turriglomina-like linings comparable also with a tube-like placen-
toid forms of Reophacidae (e.g. Subreophax scalaria Grzybows-
ki). Fig. M Klippe near Slávnické Predhorie. Thin section No.
21370,
×
150. Fig. N Klippe Krasin near Dolná Súèa. Thin section
No. 21117,
×
100.
←
118 MIÍK and SOTÁK
have been illustrated by Altiner 1991 as Meandrospiranella
sp., Pl. 13: Figs. 68), but also a tube-like placentoid
forms of Reophacidae (morphotype of Subreophax scalaria
Grzybowski).
Morphotypes of nubeculariid foraminifers
Some of these linings are irregularly shaped having an
ovate proloculus and tabular swollen chambers. They corre-
spond to the species of Nubecularia mazoviensis Bielecka &
Pozaryski (Pl. VI: Figs. B, HJ, MN). Other forms (Pl. III:
Figs. KL) are characterized by extremely elongated mostly
narrow chambers of irregular width Nubecullinella in-
fraoolithica (Terquem). A number of nubeculariids corre-
spond to the species of Nodophthalmidium jurassicum
Carozzi.
Morphotypes of uncertain linings
The linings of this morphogroup remain insufficiently
known for recognition (Pl. III: Figs. MN; Pl. V: Figs. JN;
Pl. VI: Figs. O,S,U). They are usually developed as multiloc-
ular specimens with subglobular chambers (e.g. Pl. V: Fig. J).
The glomerate-types of these linings show a certain similarity
to the saccamminid foraminifers of the genus Sorosphaera
(e.g. Pl. V: Fig. J) and Thuramminopsis (T. aff. canaliculata
Haeusler e.g. Pl. V: Fig. N). Some of them may not pertain
to foraminifera.
Conclusions
In spite of the frequent occurrences of microforaminiferal
linings in the palynological preparations no attempts to use
them for the systematical classification were made. The fora-
miniferal linings from the Callovian-Oxfordian limestones
were sometimes impregnated by Fe-oxides during early di-
agenesis, mainly in red micrites, what contributed to their
preservation and made them easily visible in thin sections.
Such impregnated organic membranes lost their plasticity, be-
came brittle and cannot be extracted by palynological meth-
ods. A set of microforaminiferal linings from the thin sections
mainly from the Callovian-Oxfordian limestones, most of
them from the neptunian dyke filling, is presented herein. The
morphological aspects of the foraminiferal linings were used
to associate them into morphogroups (foraminifers of similar
morphotypes) and form genera (foraminifers with incomplete
generic identity). Furthermore, some linings appear to be suf-
ficient for their generic as well as subgeneric classification.
Among the linings, more than 30 foraminiferal genera and
species have been classified, belonging mostly to the aggluti-
nated taxa (Textulariidae, Pseudobolivinidae, Ataxophrag-
miidae, Verneulinidae, Trochamminidae, Reophacidae,
Hormosinidae, Haplophragmoidae, Lituolidae, Nubeculari-
idae, Saccamminidae, etc.). Therefore, the linings should be
preferentially derived from the agglutinated foraminifers
(with the exception of some spirillinids and involutinids), which
secreted organic tests better than others (e.g. Jorgensen 1977).
The problem of microforaminifers consists in the lack of
agglutination in the tests, which are normally agglutinated. In
spite of this the linings show a complete morphological orga-
nization, but they ceased developing in the organic phase of
test construction. Their tiny dimensions and incomplete pres-
ervation indicates a stress sedimentary biotope of microfora-
minifers (e.g. neptunian dykes). Therefore, the abundant oc-
currence of microforaminifers in the Callovian-Oxfordian
limestones could be caused by physiochemical conditions of
sedimentation and early diagenesis. The agglutinates are
epipsammitic forms, which take from the sediment the grains
they need to build rigid tests. Considering the slow deposi-
tion rate of the Callovian-Oxfordian limestones (starvation,
strong condensation, omission surfaces, etc.), the foramini-
fers could not survive in the absence of building material for
the test agglutination. Another possibility is a total etching of
the inorganic part of the tests (submarine dissolution, bacteri-
al and algal activity, post-mortem abrasion, etc.). The abun-
dance of agglutinated foraminifers usually increases with
slower sedimentation rate (Gradstein 1983). The Czorsztyn
swell of the Pieniny Klippen Belt is thought to be upwelling
area with slow deposition and a nutrient-rich environment.
According to Tyszka & Kaminski (1995) these factors con-
Plate V: Microforaminiferal linings from the Callovian-Oxford-
ian limestones, Czorsztyn Succession, Pieniny Klippen Belt (ex-
cept Fig. G). Figs. AB Globigerina-shaped linings of trocham-
minid foraminifers (aff. Ammoglobigerina globigeriniformis (Parker
& Jones)). Fig. A Vratec-castle klippe. Thin section No. 8528,
×
200. Fig. B Klippe Kyjov-Pusté Pole. Thin section No. 20437,
×
150. Fig. C Verneulinid foraminifer stained by Fe-oxides (Duo-
taxis metula Kristan). Klippe Babina near Bohunice. Thin section
No. 12497,
×
120. Fig. D Trochamminid foraminifer Trocham-
mina globoconica Tyszka & Kaminski, Klippe Krasin near Dolná
Súèa. Thin section No. 21098,
×
160. Figs. EH Morphotypes of
spirillinid and ammodiscid foraminifers. Fig. E Glomospirella
aff. otorica Romanova, neptunian dyke filling, klippe Babina near
Bohunice. Thin section No. 12479,
×
150. Fig. F Arenoturrispir-
illina sp., neptunian dyke filling, klippe Kyjov-Pusté Pole. Thin sec-
tion No. 17181,
×
100. Fig. G Ammodiscus varians Kaptarenko-
Chernousova, pebble from the Paleocene Proè Conglomerate of the
Klippen Belt, Beòatina-II-108. Thin section No. 17168,
×
185. Fig. H
Spirillina elongata Bielecka & Pozaryski, neptunian dyke filling,
Vratec-castle klippe. Thin section No. 21091,
×
100. Fig. I Mor-
photype of bolivinid foraminifer. Sivá Skala klippe near Èervený
Kameò. Thin section No. 9588,
×
120. Fig. J Morphotype of sac-
camminid foraminifer (aff. Sorosphaera sp.). Klippe Drieòová near
Krivoklát. Thin section No. 10792,
×
185. Figs. KM Uncertain for-
aminiferal linings. Fig. K Sivá Skala klippe near Èervený Kameò.
Thin section No. 9611,
×
100. Fig. L Klippe Monèeková near Èer-
vený Kameò. Thin section No. 10343,
×
185. Fig. M Klippe Ky-
jov-Pusté Pole. Thin section No. 21101,
×
150. Fig. N Morpho-
type of saccamminid foraminifer (aff. Thuramminopsis canaliculata
Haeusler). Klippe Krasin near Dolná Súèa. Thin section No. 21098,
×
150. Fig. O Morphotype of textularid foraminifer (aff. Haghi-
mashella? sp.). Vratec-castle klippe. Thin section No. 6501,
×
150.
Fig. P Morphotype of ammodiscid foraminifer (aff. Glomospirel-
la reata (Eicher)). Neptunian dyke filling, klippe Babina near Bohu-
nice. Thin section No. 12385,
×
76.
→
PLATE V 119
120 PLATE VI
PLATE VII 121
122 MIÍK and SOTÁK
trolled the abundance of agglutinated foraminifers and other
benthics in the Czorsztyn Succession.
The organic linings of the agglutinated foraminifers show
an ability to be better protected and more resistant, and thus,
quite often preserved after death (Brönnimann & Whittaker
1988). Therefore, the well-preserved linings give the possibil-
ity to recognize their systematic position. This approach
could change the opinion about stratigraphic unimportance of
Plate VII: Microforaminiferal linings (basale membranes) ex-
tracted by the palynological procedures (material of V. Sitár) and
in the thin sections of silicites (well preserved linings without the
Fe-oxides impregnation). Figs. AB, KM Morphotypes of
haplophragmoid foraminifers (aff. Haplophragmoides concavus
(Chapman)). Figs. AB Linings extracted from the dolomites
(Hauptdolomite, Norian), basement of the Slovak part of the Neo-
gene Vienna Basin, borehole Kuklov-3, 2953 m,
×
500. Figs. KL
In a chert nodule from the Muráò Limestone, Upper Hauterivian-
Lower Barremian, Mt. Muráò, Belanské Tatry Mts. Thin section No.
12527,
×
150. Fig. M In a chert nodule from the Upper Tithonian
limestones, Pieniny Succession, Myjava. Thin section No. 19060,
×
200. Figs. CE Morphotype of bolivinid foraminifers (aff.
Pseudobolivina variana Eicher). Linings extracted from the dolo-
mites (Hauptdolomite, Norian), basement of the Slovak part of the
Neogene Vienna Basin, borehole Kuklov-3, 2953 m,
×
500. Figs. F
G Morphotype of haplophragmoid foraminifers (aff. Haplo-
phragmoides globigerinoides (Haeusler)). Linings extracted from
the dolomites (Hauptdolomite, Norian), basement of the Slovak part
of the Neogene Vienna Basin, borehole Kuklov-3, 2953 m,
×
500.
Figs. HJ Morphotype of trochamminid foraminifers (aff. Tro-
chammina inflata (Montagu)). Linings extracted from the Supra-
Posidonia Beds (Podzamcze Marl Fm.), Bajocian, Kysuca Succes-
sion of the Pieniny Klippen Belt. Radola, Kysuca Mts.,
×
500.
Plate VI: Microforaminiferal linings, naturally stained by Fe-ox-
ides, from the Callovian-Oxfordian limestones, Czorsztyn Suc-
cession, Pieniny Klippen Belt (except Figs. H,J,L). Figs. A, CG,
KL Morphotype of lituolid foraminifers belonging to the genus
Recurvoides. Fig. A Klippe Sivá Skala near Èervený Kameò.
Thin section No. 21194,
×
100. Fig. C Vratec-castle klippe. Thin
section No. 7010,
×
200. Fig. D Klippe Babina near Bohunice.
Thin section No. 21288,
×
150. Fig. E Klippe Sivá Skala near Èer-
vený Kameò,
×
100. Fig. F Neptunian dyke filling, Vratec-castle
klippe. Thin section No. 6604,
×
120. Fig. G Klippe Sivá Skala
near Èervený Kameò. Thin section No. 21193,
×
60. Fig. K
Klippe Kyjov-Pusté Pole. Thin section No. 17187,
×
100. Fig. L
Middle Liassic limestone, pebble from the Paleocene Proè Con-
glomerate, Proè. Thin section No. 16016,
×
40. Figs. B, HJ, MN
Morphotype of nubeculariid foraminifers Nubecularia mazo-
viensis Bielecka & Pozaryski. Fig. B Klippe near Krivoklát. Thin
section No. 12227,
×
150. Fig. H Red nodular Callovian-Oxford-
ian limestone of the Manín Succession, Manín Gorge. Thin section
No. 5895,
×
150. Fig. I Klippe in the Boleová Valley. Thin sec-
tion No. 19343,
×
160. Fig. J The same as Fig. H,
×
150. Fig. M
Neptunian dyke filling, klippe Babina near Bohunice. Thin section
No. 11441,
×
185. Fig. N The same. Thin section No. 12385,
×
230. Figs. OU Linings of lituolid and saccamminid foramini-
fers. Fig. O Neptunian dyke filling, klippe Babina near Bohunice.
Thin section No. 12479,
×
230. Fig. P The same. Thin section No.
12479,
×
230. Fig. R The same. Thin section No. 11441,
×
185.
Fig. S The same. Thin section No. 21288,
×
150. Fig. T Vr-
atec-castle klippe. Thin section No. 8640,
×
200. Fig. U Klippe
Krasin near Dolná Súèa. Thin section No. 21116,
×
150.
microforaminifers as was the case with the numerous other
groups of microfossils (e.g. mostly Middle Jurassic species
Trochammina aff. squamata Parker & Jones, Dentalinop-
sis aff. oolithica (Terq.), Bolivina aff. rhumbleri Franke, Tro-
cholina gracilis Blau, Pseudobolivina aff. variana Eicher,
Semiinvoluta violae Blau, Belorussiella aff. bolivinaeformis
Akimets, Ammodiscus aff. varians Kaptarenko-Chernousova,
etc.; mostly Oxfordian-Kimmeridgian species Bigenerina
jurassica (Haeusler), Haghimashella arcuata (Haeusler),
Nubecullinella infraoolithica (Terquem), Nubecularia mazo-
viensis Bielecka & Pozaryski, Ammoglobigerina globigerini-
formis (Parker & Jones), Spirillina elongata Bielecka &
Pozaryski, Glomospirella otorica Romanova, Verneulinoides
aff. minuta Said & Barakat, Thuramminopsis aff. canalicula-
ta Haeusler, Trochammina aff. pulchra Ziegler, etc.).
Acknowledgement: The authors are indebted to Dr. Stan
Stancliffe, Imperial Oil Resources Ltd., Research Center,
Calgary and Dr. Joachim Blau, Muséum dhistoire naturelle,
Genève for substancial improvements of the manuscript. The
paper contributes to the research projects Vega No 4076 and
4077 financed by the Slovak Academy of Sciences.
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