Jurassic (Bathonian–Oxfordian) planktic foraminifera from the
epicontinental strata of the Polish Basin have been investigated. The
palaeoecology, palaeobiogeography, and biostratigraphical potential of the
recorded taxa are discussed. Four species are recorded: Conoglobigerina helvetojurassica (Haeusler, 1881),
Globuligerina balakhmatovae (Morozova, 1961), G. bathoniana (Pazdrowa, 1969), and G. oxfordiana (Grigelis, 1958). This assemblage is
probably the most diverse of those described to date from the epicontinental
areas of Europe. The recorded taxa are thought to represent three different
ecological morphotypes. The clear relationship between
transgressive–regressive facies and the palaeobiogeography of the recorded
planktic foraminifera indicates a morphotype-related depth–distribution
pattern in which small, simple, globular-chambered morphotypes occupied
shallow waters whereas slightly larger, more complex forms, or those with
hemispherical chambers, inhabited deeper and more open-water environments.
Introduction
Foraminifera constitute an unique group of marine protists that are used
extensively as biostratigraphical and palaeoenvironmental proxies in the
Earth sciences. Planktic foraminifera are especially useful due to their
very wide, near-global, distribution (Masters, 1977; Hart, 1999; Premoli
Silva and Sliter, 1999; Gradstein et al., 2017a). The adaptation to a
planktic lifestyle took place relatively late in the history of the
foraminifera, as benthic forms may have appeared in the Neoproterozoic
(Pawlowski et al., 2003). It is suggested that planktic taxa appeared in the
late Early Jurassic (Toarcian, ∼ 180 Ma) following work by
Wernli (1988, 1995), Simmons et al. (1997), Hart et al. (2003), Hudson et al. (2009), Leckie (2009), Gradstein (2017), and Gradstein et al. (2017a, b).
The origin of planktic foraminifera, from benthic ancestors, is a key
question in the evolutionary history of the foraminifera and in biological oceanography in general.
Shelf environments, probably close to continental margins, were the
preferred habitat for the first appearance of primitive planktic
foraminifera (Gordon, 1970; Gradstein et al., 2017b). Because the area of
extra-Carpathian Poland was covered during the Middle and Late Jurassic by a
relatively shallow, epicontinental, sea that extended from the northern part
of the Tethys (northern Tethyan shelf), it appears to be a favourable
ecological niche for planktic foraminifera and contains an important fossil record (Olszewska
and Wieczorek, 1988; Wierzbowski et al., 2009) in both the deeper waters of
the Inner and Outer Carpathians and the shallower waters of the foreland
region of the Polish Basin (north of Kraków). What is more, the rather
low degree of diagenetic modification, and the high content of clay minerals
in the sediments of the Polish Basin, enabled the preservation of
aragonite foraminiferal tests. These mudstones also allow specimens to be extracted with little difficulty. This procedure is essential for morphological analysis
and taxonomic identification of the planktic taxa, as compared to the thin
section studies of Görög (1994) and Wernli and Görög (2000).
Wernli and Görög (1999) also resorted to acid reductions of
limestones from Hungary in order to study their material in
3-D. Despite these uniquely favourable research conditions in
the Polish Basin, Jurassic planktic foraminifera have been described by very
few authors, mostly during the 1960s and 1970s (Bielecka, 1960; Bielecka and
Styk, 1967; Pazdrowa, 1969; Fuchs, 1973; Olszewska and Wieczorek, 1988). The
aim of our research was to summarize the present knowledge of the planktic
foraminifera from the studied area as well as to analyse newly collected
material. Therefore, our research objectives were to understand the
taxonomy, evolution, and palaeoecology of the Jurassic planktic foraminifera
and demonstrate their utility in Late Jurassic stratigraphy and
palaeoenvironmental interpretations.
Before discussion of our procedures it is important to record that both
Oberhauser (1960) and Fuchs (1967, 1970) were interested in the origins of
the planktic foraminifera and, in major investigations of Triassic and Lower
Jurassic successions in Austria and northern Italy, erected a suite of both
genera and species that were based on washed residues that provided 3-D
material. Malcolm B. Hart, with the assistance of Holger Gebhardt, studied the Oberhauser
and Fuchs material in the collections of the Geological Survey of Austria
(including all of the type slides of the new taxa). This investigation
concluded that, almost certainly, none of the taxa were planktic, and this
was the same conclusion as that of Simmons et al. (1997). There are some taxa in the
collection (Oberhauserella and Praegubkinella) that look to be very close to developing a planktic
morphotype but may still be benthic in palaeoecology. Also in the
collections are the taxa erected by Fuchs (1973) based on material collected
by Manfred E. Schmid (Geological Survey of Austria) during the 10th
European Micropalaeontological Colloquium. These meetings were held on a
regular basis for many years and allowed “invited” geoscientists to visit
key localities in Europe and collect, under local guidance, material for
their institutions. The samples for the study by Fuchs (1973) appear to have
come from the upper Callovian and lowermost Oxfordian of the “Kalksteinbruch
von Wiek” near Ogrodzieniec. Both of the samples used by Fuchs (1973) – from
beds 21 and 26 – are recorded as coming from the glauconitic marls that are
present at the Callovian–Oxfordian boundary. In his paper, Fuchs (1973, p. 462, 474) describes his specimens as being “glauconitkern”, an internal cast
composed of the Fe-rich clay mineral glauconite. This preservation was also
noted by Loeblich and Tappan (1987) in their discussion of some of Fuchs'
new genera. Based on glauconitic steinkerns and with no test material or
apertures preserved, all of the new taxa erected by Fuchs (1973) are, almost
certainly, invalid
The full significance of this style of preservation only became apparent
when Plymouth-based PhD student Wendy Hudson attended the 7th
International Congree on the Jurassic System in September 2006. The field
excursion guidebook (Matyja and Wierzbowski, 2006) illustrates (op. cit., Fig. B1.8) an
excavated section in the Ogrodzieniec Quarry in which the Ogrodzieniec
Glauconitic Marls Formation (of latest Callovian and earliest Oxfordian age)
is illustrated. Samples from this temporary exposure were collected,
processed in Plymouth, and used by Hudson (2007) in her PhD thesis. Both
samples contain almost no residue (when washed on a 63 µm sieve) aside
from grains of glauconite, a very high proportion of which are the internal
casts of both benthic – but mainly planktic – foraminifera. A study of the
Fuchs (1973) collection in Vienna confirmed that the material is probably
identical and that the samples collected by Manfred E. Schmid are from the same
formation.
This comparison did resolve one problem with the illustrations displayed
in Fuchs (1973). Despite being a good likeness of the material, Malcolm B. Hart could not understand the “double” lines in many of the drawings which, in
places, are almost suggestive of marginal keels. Many of the specimens in
our material have a different colour (dark green) of glauconite within the
chambers and a slightly paler green where septa or chamber edges would have
been located.
Glauconitic steinkerns of Globuligerina sp. cf. G. oxfordiana from the Callovian–Oxfordian
boundary section, Ogrodzieniec, Poland. All scale bars represent 50 µm except 3
and 8 in which scale bars are 100 µm.
On a return visit to the Ogrodzieniec Quarry in 2011, with Jarosław Tyszka
and Eiichi Setoyama, the temporary exposure created for the Jurassic
symposium had been infilled and the Glauconitic Marls Formation was only
exposed in a track-side area (50∘46′46.5′′ N, 19∘51′86.2′′ E) with limited stratigraphical thickness visible. Samples from
this very small section yielded the same type of residue with only
glauconitic steinkerns present when washed on a 63 µm sieve. This
material is exceptionally rich in planktic foraminifera with benthic
foraminifera represented by only 10 %–15 % of the assemblage (Figs. 1, 2).
Glauconitic steinkerns of (1–8) Globuligerina sp. cf. G. oxfordiana and (9–11)
Conoglobigerina helvetojurassica from the Callovian–Oxfordian boundary section, Ogrodzieniec, Poland. All
scale bars represent 50 µm except 3, 4, 5, 6, and 11 in which scale bars are 100 µm.
Material and methods
In total, 43 samples were examined for foraminiferal studies. The samples
came from the Wrzosowa (6), Kawodrza (3), Stare Gliny (19), Ogrodzieniec
(4), Gnaszyn (7), Bolęcin (1), Czatkowice (1), Gorenice (1), and Podłęże (1) quarries located in the Polish Jura (Fig. 3), which,
combined, expose strata from the Morrisi Zone (middle Bathonian, Middle
Jurassic) to the Bifurcatus Zone (upper Oxfordian, Upper Jurassic). We have
sampled all available stratigraphical divisions (ammonite zones).
Specifically, the Morrisi Zone (middle Bathonian) was sampled in the
Kawodrza (50∘48′09.8′′ N, 19∘04′01.2′′ E) and Gnaszyn
(50∘48′05.5′′ N, 19∘02′42.3′′ E) clay pits. These locations
display a rather homogenous succession of black clays, with the sporadic
occurrence of siderite nodule horizons (Matyja and Wierzbowski, 2003; Matyja
et al., 2006b; Leonowicz, 2012; Parent and Zatoń, 2016).
Geological pre-Cenozoic sketch map of the Polish Jura and
adjoining areas (after Wierzbowski et al., 2009) showing the location of
sections studied and previously described in the literature: Chrzanów
(Bielecka, 1960), Ogrodzieniec (Bielecka and Styk, 1967; Pazdrowa, 1969), and
Gnaszyn (Smoleń, 2012).
Upper Bathonian and lower to middle and middle Callovian deposits that
developed as clays and glauconitic marls respectively were sampled in
Ogrodzieniec (50∘27′54.6′′ N, 19∘31′17.2′′ E) by Dembicz in
2006 (Dembicz and Praszkier, 2003; Matyja and Głowniak, 2003; Barski et
al., 2004; Dembicz et al., 2006; Wierzbowski et al., 2009; Leonowicz, 2013).
The Glauconitic Marls Formation (uppermost Callovian to lowermost Oxfordian)
was, as reported above, sampled by Hudson in 2006 and by Malcolm B. Hart in 2011. At the
present time, the lower part of the outcrop is unavailable.
An incomplete succession from the Jason Zone (middle Callovian) to the
Bifurcatus Zone (middle to upper Oxfordian) was sampled exclusively in the
Stare Gliny Quarry (50∘21′00.8′′ N, 19∘35′29.2′′ E)
(Wierzbowski et al., 2009, 2021). This important Jurassic section is about 21 m thick and includes middle to upper Callovian marl–limestone rhythmites (10
samples; 9 m) and lower to upper Oxfordian clay strata (9 samples;
12 m).
Jurassic strata in the Wrzosowa Quarry (50∘45′25.4′′ N,
19∘08′51.9′′ E) are represented by a succession, approximately 5 m
thick, of Cordatum Zone (lower Oxfordian) marl–limestone rhythmites with the
dominant limestone of the Jasna Góra beds, yielding numerous sponges,
belemnites, ammonites, and brachiopods (Trammer, 1985; Matyja et al., 2006a;
Wierzbowski et al., 2009; Głowniak, 2012). Some isolated samples of
Callovian and Oxfordian strata were additionally collected from little-known
sections such as Bolęcin (lower Callovian) (Różycki, 1953),
Czatkowice (Lamberti Zone, upper Callovian) (Dayczak-Calikowska and Kopik,
1973a; Salata, 2013), Gorenice (lower Oxfordian) (Różycki, 1953),
and Podłęże (middle Oxfordian) (Giżejewska and Wieczorek,
1977; Jurkowska and Kołodziej, 2013).
All of the collected samples are characterized by high clay content, which
provides excellent conditions for fossil preservation, enabling the
acquisition of isolated free specimens from the sediments (Hart et al.,
2019). The use of 3-D specimens instead of thin sections (see the
work of Hudson et al., 2009, on material from the Inner Carpathians)
afforded us improved data for detailed morphological analysis. Selectively
collected clay-rich samples were disintegrated using Glauber's salt
(Na2SO4×10H2O) or detergent (Witwicka et al.,
1958). The material was then washed in an ultrasonic cleaner and sieved
using two sieves: 0.071 mm (lower) and 0.6 mm (upper). Samples that
required additional cleaning were treated with liquid nitrogen (Remin et
al., 2012). Foraminifera were manually picked from the residue and studied
with the use of a Nikon SMZ18 stereoscopic microscope. Taxonomic
observations and images were generated using a Zeiss Sigma VP scanning
electron microscope at the Faculty of Geology, University of Warsaw.
Results
In general, planktic foraminifera in the samples were abundant, if not very
numerous, and variably preserved. In all of the samples from Wrzosowa and
some from Gnaszyn (G1252, G1264, G1318), Kawodrza (K1), Ogrodzieniec (O1, O2, O3), and Stare
Gliny (ST6B, ST6F, ST6H, ST18, ST0, ST60, ST560), planktic forms were
absent. The scarcity of foraminifera was almost certainly caused by the
dissolution of their aragonitic tests, as all of the aragonitic benthic
foraminifera had also been lost and the foraminiferal assemblages were
represented only by benthic foraminifera with calcitic wall structures.
Planktic foraminifera were recorded in the other studied samples, although
their state of preservation was variable. Rather poorly preserved specimens
with obscure morphological features, such as ornamentation or apertures
(some of the specimens were preserved as glauconitic internal moulds, as
described above in material from Ogrodzieniec) were collected from the
Callovian of Stare Gliny (samples 2, 4, 16), Czatkowice, Gorenice, and Podłęże, whereas very well-preserved specimens were obtained mostly from
samples that came from Gnaszyn (sample 1330), Kawodrza (samples 2, 3), and
Stare Gliny (samples 140, 320, 500).
SEM images of studied planktic foraminifera. Globuligerina balakhmatovae (Morozova, 1961)
specimens: (1a–b) Stare Gliny, sample 140, MWGUW ZI/67/MG4.65 and (2a–b)
Stare Gliny, sample 140 MWGUW ZI/67/ZD64ST14. Globuligerina bathoniana (Pazdrowa, 1969) specimens:
(3, 4) Gnaszyn, sample 1324, MWGUW ZI/67/MG1.01 and (5) Stare Gliny, sample
140, MWGUW ZI/67/ZD64ST02. Globuligerina oxfordiana (Grigelis, 1958) specimens: (6) Podłęże, sample 1, MWGUW ZI/67/MG1.42; (7) Podłęże, sample 1,
MWGUW ZI/67/MG1.20; (8) Stare Gliny, sample 6, MWGUW ZI/67/MG4.62; and (9)
Stare Gliny, sample 140, MWGUW ZI/67/ZD64ST03. Conoglobigerina helvetojurassica (Haeusler, 1881) specimens:
(10) Stare Gliny, sample 380, MWGUW ZI/67/MG3.12; (11) Stare Gliny, sample
120, MWGUW ZI/67/MG4.57; (12) Stare Gliny, sample 140, MWGUW ZI/67/MG4.07.
Characteristic wall ornamentation (13-14) of genus: (13) Globuligerina, Gnaszyn, sample
1312, MWGUW ZI/67/MG1.02 and (14) Conoglobigerina, Stare Gliny, sample 140, MWGUW
ZI/67/MG4.49. All scale bars represent 100 µm.
In our material, four species from two genera were recorded:
Conoglobigerina helvetojurassica (Haeusler, 1881), Globuligerina balakhmatovae (Morozova, 1961), Globuligerina bathoniana (Pazdrowa, 1969), and Globuligerina oxfordiana (Grigelis, 1958)
(Fig. 4). C. helvetojurassica is characterized by a low to medium-high trochospiral large test;
the mean diameter measured on 10 randomly selected specimens was 260 µm.
The almost spherical chambers are much larger in the last whorl than those
in the previous whorl, and they sit close to each other. Well-preserved
pseudomuricae are visible on the surface of the test.
G. balakhmatovae has a much smaller and low trochospiral test with oval, almost petaloid
chambers. The chamber flattening, commonly observed in the studied
specimens, is apparently a primary feature and is not produced during
taphonomy. The main diagnostic features of G. bathoniana are a high trochospiral test, spherical
chambers, and a test surface almost covered by pustulose ornamentation. The
mean diameter of G. bathoniana, measured on 10 randomly selected specimens, was 160 µm. Similarly, G. oxfordiana is characterized by pustulose ornamentation. The coiling of
the test is, however, that of a relatively low trochospiral. The last
chamber is significantly larger than earlier ones and is often more
elongated. The mean size of 10 randomly selected specimens of G. oxfordiana was 150 µm.
The most common taxa in our material were G. bathoniana and G. oxfordiana.
G. bathoniana was the most numerous in Bathonian strata, especially in the Morrisi Zone
(middle Bathonian) of Gnaszyn and Kawodrza, and in the Callovian and
Oxfordian sediments of Bolęcin, Czatkowice, Ogrodzieniec, Podłęże, and Stare Gliny. A relatively similar stratigraphical
distribution was observed for G. oxfordiana, which was a dominant component of the
Czatkowice, Podłęże, and Stare Gliny assemblages but rare in the
Bathonian samples from Gnaszyn and Kawodrza. Specimens of C. helvetojurassica were recorded
exclusively in the upper Oxfordian (Bifurcatus Zone) sediments from Stare
Gliny (samples 140, 200, 380). Well-preserved specimens retained the
characteristic pseudomuricae ornamentation, which is a diagnostic feature of
the genus Conoglobigerina (Simmons et al., 1997; Gradstein et al., 2017a). Specimens of G. balakhmatovae
were found in four samples from Kawodrza (Morrisi Zone, middle Bathonian)
and from the Bifurcatus Zone (of Stare Gliny (upper Oxfordian) samples 140,
320, 380). No hemispherical forms were recorded from any Callovian
(Jason–Lamberti zone) sediments.
DiscussionSpecies variability of Jurassic planktonic foraminifera in
extra-Carpathian Poland
Beginning in the second half of the 20th century, Jurassic planktic
foraminifera have been investigated worldwide (Gradstein et al., 2017a),
although few investigations have been undertaken in extra-Carpathian Poland.
The first records derive from the Polish Jura where, in the late 1960s,
Globuligerina sp. cf. G. oxfordiana and Globuligerina sp. cf. G. helvetojurassica (Bielecka and Styk, 1967) were found in middle–upper
Callovian (Jason–Lamberti zones) and Mariae Zone (lower Oxfordian) strata
of Ogrodzieniec. In 1969, Olga Pazdrowa described G. bathoniana from the Morrisi Zone
(middle Bathonian) exposed in this quarry. It should be remembered, however,
that her work was carried out in 1959 as that is the date written (by her)
on the slides in the collections of the institute in Kraków. Several
years later, Fuchs (1973) published his description of planktic foraminifera
from the Polish Jura, identifying many new species. As all of this
material is illustrated by line drawings of moulds, without original test
walls, ornamentation, and apertures, none of these new taxa are valid
(Loeblich and Tappan, 1987; Hudson, 2009; Gradstein et al., 2017a); see
earlier discussion.
Subsequently, several sites from extra-Carpathian Poland were studied in
terms of Jurassic planktic foraminifera (Kraców region, by Olszewska and
Wieczorek, 1988; Ogrodzieniec, by Hudson, 2009, and Hart et al., 2012; and Gnaszyn, by
von Hillebrandt, 2012, Smoleń, 2012, and Kendall et al., 2020). Moreover,
planktic foraminifera were recorded in microfossil assemblages from
boreholes in northern Poland (Chrzanów, by Bielecka, 1960; Kcynia, by
Bielecka and Styk, 1964; Pasłęk, by Bielecka and Styk, 1966;
Bartoszyce, by Bielecka and Styk, 1966, and Bielecka, 1974; Grzybnica, by
Bielecka and Styk, 1981; and Wolin, by Bielecka and Styk, 1981) where, apart
from G. bathoniana from Wolin (Fig. 5),
G. oxfordiana was the only recorded planktic taxon.
Palaeogeographical maps of the (a) middle Oxfordian, (b) middle
Callovian, and (c) middle Bathonian of central Europe, with the distribution of
planktonic foraminifera based on data in the subject literature (Bielecka,
1960, 1974; Bielecka and Styk, 1964, 1966, 1981) and recent studies.
To summarize, planktic foraminifera from extra-Carpathian Poland are rather
scarce and require additional investigation. In fact, to date, only two
species of planktic foraminifera, G. bathoniana and G. oxfordiana, have been described with certainly
from this region. It is worth mentioning that Bielecka and Styk (1967, 1981)
recorded C. helvetojurassica in this area of Poland, although it is clearly indicated in their
papers that this record is uncertain. There is only one very poor
illustration of this species included in the work of Bielecka and Styk
(1981), and this is probably inadequate for a reliable taxonomic
identification.
G. oxfordiana is commonly regarded as a cosmopolitan Jurassic planktic form (Hudson et
al., 2009), but this is probably the result of many authors naming any
mid-Jurassic planktic form as this species. The newly studied Oxfordian
assemblage from Stare Gliny is quite species-rich and probably one of the
most diverse in the previously described area of epicontinental
extra-Carpathian Poland and Europe in general. In this regard it is similar
to the rather diverse assemblage seen as glauconitic steinkerns in the
material from Ogrodzieniec described here. Specifically, this assemblage is
composed of common G. bathoniana and
G. oxfordiana but also includes the typical Tethyan C. helvetojurassica and
G. balakhmatovae. These four taxa represent three different morphotypes: a small globular-chambered morphotype, represented by G. bathoniana and G. oxfordiana; the hemispherically chambered
G. balakhmatovae; and the often larger and more complex C. helvetojurassica.
Biostratigraphical remarks
Gradstein et al. (2017a, b) show that the rate of speciation and
evolutionary diversification of the first planktonic foraminifera was rather
slow, confirming earlier suggestions by Simmons et al. (1997), Premoli Silva
and Sliter (1999), Hart (1999), and Hart et al. (2002). Consequently, even
the later Jurassic forms were characterized by a low level of morphological
and taxonomic variability, which is, unfortunately, a serious disadvantage
for stratigraphical studies. Additionally, the low preservation potential of
the aragonitic tests of Jurassic planktic foraminifera very often results in
their poor preservation or complete absence from the fossil record.
Not many species of Jurassic foraminifera are unquestionably considered as
planktic. Gradstein et al. (2017a) described only 10 different species:
Globuligerina dagestanica (Morozova), G. avariformis (Kasimova), G. balakhmatovae (Morozova), G. oxfordiana (Grigelis), G. bathoniana (Pazdrowa), G. jurassica (Hofman),
G. oxfordiana (Grigelis) calloviensis Kuznetsova, G. tojeiraensis Gradstein, Conoglobigerina helvetojurassica (Haeusler), C. grigelisi Gradstein, and C. gulekhensis
(Gorbachik and Poroshina). Recently, however, Apthorpe (2020) described
three new planktic taxa: Globuligerina bathoniana australiana n. ssp., Globuligerina altissapertura n. sp., and Mermaidogerina loopae n. gen. n. sp. from the
Bajocian of the southern Tethys (north-west Australia). These records, from dredge
samples, lack precise dating, as the Jurassic material is entombed in
Pleistocene sediments, but the occurrence – so far away from the otherwise
European and North Atlantic Ocean locations – creates problems for
palaeogeographical reconstructions (see Hudson et al., 2009). This
assemblage is, perhaps, more reminiscent of the Bathonian–Oxfordian
interval than the Bajocian.
This rather low level of taxonomic diversity, the scattered occurrence and
rather poor taphonomic potential of Jurassic planktics, and the lack of
extensive sampling have resulted in a rather low level of resolution for
standard biostratigraphical zonation (see Ogg et al., 2016). The most
recent and exhaustive zonation of Jurassic planktonic foraminifera presented
by Gradstein et al. (2017b) includes six zones (from J1 to J6).
Unfortunately, the total ranges of specific zones have been established only
tentatively and are still uncertain; no links to ammonite zones have been
assigned. Nonetheless, Jurassic planktic foraminifera possess significant
interregional correlative potential, as described from a wide
palaeogeographical area including the eastern part of North America, Europe, the
North Atlantic Ocean, the Gulf of Mexico, North Africa, the Middle East, and
Australia. Additional investigations are, therefore, definitely needed in
order to create a robust biostratigraphical zonation correlated with other
groups of index fossils (especially calcareous nannofossils and ammonites).
The stratigraphical distribution of the studied taxa from the Polish Basin
closely matches the biostratigraphical scheme of Gradstein et al. (2017b).
In the studied area, the earliest forms recorded are G. bathoniana and G. balakhmatovae, which occurred
in the Morrisi Zone (middle Bathonian) of Gnaszyn (see also Pazdrowa, 1969;
Bielecka and Styk, 1981; Hart et al., 2012; Smoleń, 2012; Kendall et
al., 2020) and Kawodrza (Fig. 6). G. bathoniana occurs frequently in the Morrisi Zone
(middle Bathonian) of the ore-bearing Czestochowa clays in which it was
originally described (Pazdrowa, 1969). Both G. bathoniana and G. balakhmatovae are known from the late
Bathonian of other locations on the margins of the Tethys, namely, Grand
Banks in Eastern Canada (Gradstein, 1979; Stam, 1986) and the lower
Bathonian (Zigzag Zone) of central Dagestan (Morozova and Moskalenko, 1961).
In general, G. bathoniana appeared slightly earlier than G. balakhmatovae and is characterized by a wider
stratigraphical range (Gradstein et al., 2017b) encompassing the interval
from the early Bajocian to the late Tithonian, whereas the latter appeared
from the middle Bajocian to the end of the Kimmeridgian (Fig. 5). Whereas
G. bathoniana was constantly recorded from the Bathonian up to the Oxfordian in the
studied area, G. balakhmatovae was missing from the Callovian samples (Fig. 6).
Chronostratigraphy, lithological columns, and ranges of recorded
species of planktonic foraminifera in the studied sections, including a
comparison with their total Tethyan ranges (Gradstein et al., 2017b).
The general stratigraphical range of G. oxfordiana is assessed as early Bajocian–late
Tithonian – i.e. exactly the same range as that of G. bathoniana (Gradstein et al., 2017b).
The first appearances of both taxa were established as the basis for the
Jurassic planktic foraminiferal zone J2. Its lower Bajocian and Bathonian
occurrence has been confirmed from, for example, Hungary (Humphriesianum and
Niortense zones; Wernli and Görög, 1999) and Eastern Canada (Stam,
1986) respectively. These records challenge the hypothesis that the range
of this taxon is restricted to the Oxfordian stage (Simmons et al., 1997).
In the studied area, C. helvetojurassica has been found only in the upper Oxfordian (Bifurcatus
Zone) of Stare Gliny (Fig. 6); this is the first record of this taxa in
extra-Carpathian Poland. C. helvetojurassica has been typically described from Tethyan strata,
e.g. the middle Oxfordian Transversarium Zone of Switzerland (Gradstein et al.,
2017a) and the lower Kimmeridgian Planula–Platynota zones of Portugal (Stam,
1986; Gradstein, 2017; Gradstein et al., 2017a) and France (Görög and Wernli, 2013).
In general, C. helvetojurassica ranges from the middle Oxfordian to the early Kimmeridgian; its
first appearance defines the beginning of the planktonic foraminiferal zone
J5.
The co-occurrence of the four species C. helvetojurassica, G. balakhmatovae, G. bathoniana, and G. oxfordiana in the marls of the Stare
Gliny section indicate zone J5 of Gradstein et al. (2017b) (Fig. 6). This
zone, ranging from the middle Oxfordian to the early Kimmeridgian, is
defined by the first appearance of the genus Conoglobigerina as well as the common
occurrence of typical simple and hemispherical forms such as G. balakhmatovae, G. bathoniana, and G. oxfordiana.
Regrettably, due to the wide stratigraphical ranges of Jurassic planktic forms
and the lack of characteristic events, no other zones were identified from the
remainder of the sampling material.
Distribution of Jurassic planktonic foraminifera in extra-Carpathian
Poland and their palaeoenvironmental potential
Planktic foraminifera are undoubtedly very useful palaeoenvironmental and
palaeoecological proxies due to their close relationship with test
morphology (morphotype) and with the habitat they occupied. Moreover, their
tests constitute a useful medium for recording stable (carbon and oxygen)
isotopes or ratios of trace elements such as Mg and Sr.
The geographical distribution of Holocene planktic foraminifera is restricted to
fully marine habitats from the Equator to relatively high latitudes. These
organisms occur in the water column from the surface water to several
hundred metres in the water column (Bé and Hamlin, 1967; Bé, 1977;
Schiebel and Hemleben, 2005, 2017). Thus, the union of Tethys and the
proto-Atlantic with adjacent epicontinental seas during the mid-Late
Jurassic apparently resulted in the expansion of planktic forms (Hart et
al., 2002; Hudson et al., 2009; BouDagher-Fadel, 2015). Planktic
foraminifera, having originated in the Tethys Zone, subsequently migrated
into adjacent epicontinental seas, and a brief review of their development
follows.
The earliest planktic foraminifera appear to be those described by Wernli
(1995) from the Creux de l'ours section in the area of Teysachaux
(Swiss Prealps, Fribourg, Switzerland). The mudstones from which the
planktic foraminifera were recovered are located within a few metres of the
Toarcian Oceanic Anoxic Event (OAE), an event recently re-interpreted by Ruebsam et al. (2019) as
possibly related to warming and sea level rise following a glacial (or cold)
event.
In the UK, Hylton (2000) and Hart et al. (2003) recorded a flood of inflated
Oberhauserella or Rheinholdella at the same stratigraphical level immediately above the Toarcian OAE
(Whitby Mudstone Formation). Planktic foraminifera are rare in the overlying
Aalenian (Wernli, 1988), becoming more abundant (and widespread) in the
Bajocian (Gradstein et al., 2017b). Many of the mid-Jurassic occurrences are
in “Ammonitico Rosso” facies in which ammonites and calcite “filaments” are
the only other fossils. The important questions are as follows:
Are these mid-Jurassic planktic foraminifera with simple test arrangements
surface-dwelling forms?
Do all of these Jurassic planktic foraminifera have aragonitic tests and are
they, therefore, a completely unrelated lineage to most Cretaceous taxa?
Why are Jurassic assemblages so species-poor and why did they never developed greater
diversity?
In answer to question 1, the simple “globigerine” forms, which often have only three to four
chambers in the final whorl, appear comparable to many Cretaceous taxa in
which stable isotope analysis (Wendler et al., 2013) indicates the
occupation of surface waters (above the thermocline). In a recent study of
Globuligerina spp. from Poland and Portugal, Kendall et al. (2020) – using micro computed tomography (micro-CT)
analysis – suggest that this morphology reflects short life cycles and
potentially rapid reproduction in nutrient-rich coastal environments.
In the Czorsztyn Limestone Formation (Oxfordian) of the Pieniny Klippen Belt
of Poland, Hudson et al. (2009) described thin sections full of simple,
globigerina, forms (often with a maximum of four chambers in the final whorl)
and very few associated benthic taxa. These were thought to represent
open-ocean environments in which a 99:1P:B ratio indicates planktic ooze
deposition above a Jurassic aragonite compensation depth (ACD), although the
actual water depth is impossible to estimate. The palaeoenvironmental
evidence of Hudson et al. (2009) and Hart et al. (2012), coupled with the
morphological analysis of Kendall et al. (2020), is suggestive of a
surface-water dwelling habitat for most Jurassic planktic foraminifera.
These species were, therefore, able to spread into shelf seas only during
sea level highstands (Gordon, 1970), which resulted in the minimum seawater
depth required for these organisms to thrive. It is also believed that
foraminiferal migration farther into the shelves was additionally associated
with the occasional incursion of warm currents (Gordon, 1970; Riegraf, 1987;
Hudson et al., 2009). Therefore, it is possible that planktic foraminifera
first appeared in the Polish epicontinental seas during a transgressive
interval in the late Bathonian (Morrisi Zone) (Dayczak-Calikowska and Kopik,
1973b; Leonowicz, 2016), where they were recorded (Fig. 5) in the Polish
Jura (Pazdrowa, 1969; Fuchs, 1973; Smoleń, 2012) as well as in northern
Poland (Bielecka and Styk, 1981). No planktic foraminifera have been
recorded in the Aalenian and Bajocian, as these stages are represented by
very shallow, although transgressive, facies only as the Aalenian sea
entered the Polish Basin (Dayczak-Calikowska, 1997). Until the late
Bajocian, the basin in the Polish Lowlands was very narrow (elongated in a
north-western–south-eastern direction) and had not expanded towards the north-eastern part of
Poland (Dayczak-Calikowska and Kopik, 1973b). During the late Bathonian, a
significant expansion took place and, as a result, the sea covered nearly
all of the Polish Lowlands (Dayczak-Calikowska and Kopik, 1973b; Leonowicz,
2016). The first planktic foraminifera from the area of Poland are actually
recorded from the Bathonian: G. bathoniana and G. oxfordiana (e.g. Pazdrowa, 1969; Smoleń, 2012; Hart
et al., 2012; Kendall et al., 2020).
A short-term, relatively rapid, regression occurred in the early Callovian,
particularly affecting the basin in the Polish Lowlands (Dayczak-Calikowska
and Kopik, 1973b);
G. bathoniana and G. oxfordiana have rarely been recorded subsequently, even in the southern parts of
the Polish Basin (Bielecka and Styk, 1967; Bielecka, 1960; Bielecka and
Styk, 1981). Similarly, as in their early Callovian distribution, both
species occurred sporadically in the middle and upper Callovian samples from
Stare Gliny and Ogrodzieniec. Contrastingly, during the late
Callovian–Oxfordian interval, planktic foraminifera expanded to attain
their widest distribution, described as the “foraminiferal flood” across
Europe (Hudson et al., 2009), which was apparently connected with global
oceanographical and climatic changes occurring at the transition between the
Middle and Upper Jurassic (Callovian to Oxfordian). Specifically, a marked
transgression, beginning in the late Callovian and continuing until the
Kimmeridgian (Dadlez et al., 1998), resulted in the appearance of widespread
epicontinental seas, which communicated with the Tethys covering much of Europe, as well as in climate warming.
Mediterranean ammonites began to be found instead of boreal taxa in the
Oxfordian strata of the Polish Jura (Dąbrowska, 1973), and this was simultaneous with
with the first occurrence of typical Tethyan foraminifera (C. helvetojurassica and
G. balakhmatovae) which were recorded in our studies in the Oxfordian of Stare Gliny. In
general, in the Oxfordian, planktic foraminifera (G. oxfordiana) expanded to the northern
margins of the Polish Basin (Fig. 5; Bielecka, 1974; Bielecka and Styk, 1964, 1966,
1981), as well as in the Lithuanian and Russian basins (Grigelis,
1958, 1977; Grigelis, 2016; Gradstein et al., 2017a). Excellently preserved
G. oxfordiana specimens are recorded from lower Oxfordian dark clay- and siltstones of
Lithuanian boreholes (Grigelis, 1958) and middle to late Oxfordian strata of
central Russia (Grigelis, 1977; Gradstein et al., 2017a). The abundance of
planktonic foraminifera in the Russian Sea (Makar'yev section, Volga River Basin, Russia) during the middle Oxfordian is
related to palaeogeographical changes, such as sea level highstands
associated with the opening of the pathway connecting the northern Tethys with
the Boreal realm (Colpaert et al., 2016). Moreover, it was in the Oxfordian
that planktic foraminifera expanded northwards; G. oxfordiana has been recorded from lower Oxfordian strata of Sweden (Gorbachik and
Kuznetsova, 1983).
The depth distribution and variability of Holocene planktic foraminifera is
related to morphology (Bé and Hamlin, 1967; Bé, 1977; Schiebel and
Hemleben, 2005, 2017), and this relationship has been used in the
interpretation of habitats of ancient morphotypes. The connection
between the life cycle of planktic foraminifera and depth stratification is
expressed as a depth–morphogroup model commonly applied to Cretaceous (Hart
and Bailey 1979; Hart, 1980; Caron and Homewood 1983; Leckie, 1987) and
Cenozoic (Keller, 1985) palaeoenvironmental interpretations. According to
this general model, the simple, small globigerina-like morphotypes of the
Cretaceous lived near the surface of the water, while more complex, larger,
hemispherical- or keel-chambered morphotypes preferred deeper to intermediate
habitats. Unfortunately, no depth–morphogroup model has been proposed for
Jurassic taxa. Jurassic foraminifers are probably a separate lineage than
Cretaceous–Holocene foraminifera in possessing aragonite instead of calcite
shells. The only lineage that appears to cross
the Jurassic–Cretaceous boundary is that which includes the favusellids, and
this genus extends, in relative low numbers, up into the mid-Cenomanian, with the
last species being Favusella washitensis (Carsey, 1926). There are no other lineages that appear
to develop from the favusellids. However, when we analysed the distribution
of the Jurassic planktonic foraminifera in the Polish Lowlands in relation
to the sea level curve as well as to facies distribution and the
palaeogeographical map (distance from the ocean) (Hudson et al., 2009; Wierzbowski et al., 2009; Leonowicz, 2016), the
depth–distribution pattern of Jurassic planktonic foraminifera appeared
comparable to that of the Cretaceous. Specifically, the shallow shelf facies
and the areas quite distant from the open ocean yielded only small, simple
morphotypes such as G. bathoniana and G. oxfordiana. Planktic foraminiferal assemblages composed of
only these two morphologically simple forms have been collected mostly from
the Bathonian and Callovian, as these strata were deposited in a still
relatively shallow sea. Only monospecific G. oxfordiana planktonic assemblages have been
recorded in the Oxfordian strata of the northern part of the Polish Basin,
implying that, in the Oxfordian, the northern margin of the basin was
invaded by only shallow-dwelling forms, in contrast to the southernmost
part of the basin, which represented a deeper-shelf environment, where
planktic foraminiferal assemblages were slightly more diverse and
represented by more complex and larger forms, such as G. balakhmatovae and
C. helvetojurassica. This is suggestive of a deepening of the environment towards the outer
shelf and open ocean. To summarize, assemblages composed of only simple
forms such as
G. bathoniana and G. oxfordiana were typical of rather shallow- and mid-shelf environments, and, by
analogy to Holocene counterparts, may have been characterized by short life
cycles which may have unfolded in relatively shallow waters beyond the open
ocean. The Oxfordian of the Stare Gliny outcrop, deposited in an outer-shelf
environment, closer to the Tethys (actually, Stare Gliny represents the
southernmost available Jurassic strata of the Polish epicontinental basin),
contains more diverse planktic assemblages. In addition to simple G. bathoniana and G. oxfordiana, it contains hemispherical G. balakhmatovae and the larger morphotype C. helvetojurassica. The latter was
the most evolutionarily advanced species of all those reported in this study
and may have needed a much greater water column to establish favourable
conditions for its life cycle.
Conclusions
Four species of Jurassic planktonic foraminifera, Conoglobigerina helvetojurassica (Haeusler, 1881), Globuligerina balakhmatovae (Morozova, 1961), Globuligerina bathoniana (Pazdrowa, 1969), and Globuligerina oxfordiana (Grigelis, 1958), were identified in material from
extra-Carpathian Poland. In general, the level of richness of the studied
assemblages was rather low and undifferentiated. The majority were
represented by only two common taxa,
G. bathoniana and G. oxfordiana. Nevertheless, an assemblage from the middle–upper Oxfordian clays and
marls of the Stare Gliny Quarry includes all four of the species mentioned
above, making it currently the most diverse planktic foraminiferal
assemblage of the European epicontinental Jurassic.
The stratigraphical distributions of all of the studied taxa closely match the general biostratigraphical zonation for Jurassic planktic foraminifera (Gradstein et al., 2017b).
The first planktic foraminifera from the Polish Basin, G. bathoniana and G. balakhmatovae, derive from
middle Bathonian (Morrisi Zone) transgressive deposits of the Polish Jura.
In the middle–upper Callovian (Jason–Lamberti zones) of the Polish Jura,
planktic foraminifera, represented by G. bathoniana and G. oxfordiana, are discontinuously and rather
sparsely recorded. Due to the late Callovian–Oxfordian transgression,
planktic foraminifera migrated to the northern part of the Polish area,
where they are represented mostly by monospecific assemblages of G. oxfordiana, in
contrast to the southern part, where assemblages are more diverse and
contain complex Tethyan forms.
Three separate test morphotypes within the studied Jurassic planktic
foraminifera were distinguished: a small and simple globular-chambered
morphotype represented by
G. bathoniana (the mean diameter of all the studied specimens is ca. 160 µm) and
G. oxfordiana (the mean diameter of the studied specimens is ca. 150 µm),
a hemispherically chambered morphotype by G. balakhmatovae, and a large and complex
morphotype by C. helvetojurassica (the mean diameter of all the studied specimens is ca. 260 µm). These morphotypes apparently reflect the close relationship
between the settled habitat and test morphology. The deeper the environment, the
more diverse the foraminiferal assemblages that occurred there, with a
significant share of large and complex forms. Contrastingly, small and
simple foraminifera dominated the shallow inner-shelf environments. This
distribution pattern would have some similarities to depth–distribution
models created for younger forms such as Late Cretaceous planktic
foraminifera (Leckie, 1987; Caron and Homewood, 1983). Thus, beyond doubt,
Jurassic planktic foraminifera may also be regarded as useful depth
indicators for palaeoenvironmental studies.
In general, our studies provide new insight into the palaeoecology,
evolution, and distribution of Jurassic planktic foraminifera from the
Polish Jura. However, the group still requires more investigation.
Author contributions
ZD conceived this research project. MG processed and analysed the samples.
All authors interpreted results and wrote and edited the article.
Competing interests
Malcolm B. Hart reviewed the first version of this paper and suggested substantive improvements. He then participated as a co-author when the paper was revised.
Acknowledgements
We kindly thank Krzyś Dembicz for his
invaluable help during fieldwork. Christopher W. Smart (University of Plymouth) generated Figs 1 and 2 from the SEM micrographs. Jarosław Tyszka and Eiichi Setoyama
assisted with the fieldwork in 2011.
Holger Gebhardt (Geological Survey of Austria) is thanked for providing
access and microscope facilities for work on the collections of Rudolf Oberhauser and Werner Fuchs.
Financial support
This research has been supported by the National Science Centre, Poland (grant no. UMO-2017/27/B/ST10/00687).
Review statement
This paper was edited by Laia Alegret and reviewed by Malcolm B. Hart, Mike Simmons, and one anonymous referee.
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