FRFossil RecordFRFoss. Rec.2193-0074Copernicus PublicationsGöttingen, Germany10.5194/fr-20-9-2016The Late Devonian placoderm Aspidichthys Newberry, 1873 from the Holy Cross Mountains, PolandDworczakPatrycja G.SzrekPiotrpiotr.szrek@pgi.gov.plInstitute of Geology, Institute of Geology, Adam Mickiewicz University, 16 Maków Polnych Street, 61-606 Poznań, PolandPolish Geological Institute – National Research Institute, 4 Rakowiecka Street, Warsaw, PolandPiotr Szrek (piotr.szrek@pgi.gov.pl)21November201620191923June20162September20167September2016This work is licensed under a Creative Commons Attribution 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by/3.0/This article is available from https://fr.copernicus.org/articles/20/9/2016/fr-20-9-2016.htmlThe full text article is available as a PDF file from https://fr.copernicus.org/articles/20/9/2016/fr-20-9-2016.pdf
The reported placoderm remains from the Holy Cross Mountains, Poland, belong
to the species Aspidichthys ingens Koenen, 1883. This study focuses
on the material described in the past from Wietrznia Quarry but the new
specimens were also collected form Kowala and Płucki localities. All
specimens presented here differ from others described in the past, mainly
by having smaller tubercles and possessing a distinctive crest in the anterior
median part of the median dorsal plate, but did not allow the erection of a
new taxon. Based on conodont assemblage, all specimens are attributed to a
Frasnian age. Aspidichthys occurs in the middle to the upper part
of the Frasnian (Palmatolepis punctata to P. linguiformis
conodont zone) and is not found in the Famennian boundary. Described
specimens also occur in a stepwise deepening environment caused
by local synsedimentary tectonics.
Introduction
The genus Aspidichthys has an uncertain systematic position and is
traditionally placed within the Arthrodira and closely related to the
holonematids (e.g. Kulczycki, 1957; Schultze, 1973; Denison, 1978). It
comprises large and medium-size placoderms with a distinctive elongated
median dorsal plate, and well-marked large tubercles cover dermal bones. The
tubercles are closely spaced and irregularly arranged. They may form rows
paralleling the plate outline, usually at the margins. Plates are relatively
thick (reach several centimetres). Due to these features, it is easy to
recognize even partially preserved specimens of Aspidichthys,
which has noticed very often by many authors, in several areas of the former
Laurussia and Gondwana. Remains of Aspidichthys are known from the USA
(Newberry, 1873; Sinclair and Walker, 1956; Miles, 1965); north-eastern Iran
(Schultze, 1973); Morocco (e.g. Rücklin, 2010), Bad Wildungen, Germany
(e.g. Gross, 1932); and the Holy Cross Mountains, Poland (Kulczycki, 1957;
Ivanov and Ginter, 1997; Janiszewska et al., 2007; Szrek, 2006, 2007, 2008;
Szrek and Ginter, 2007, 2008).
The main goal of this paper is to describe placoderms previously assigned to
the genus Aspidichthys from the Upper Devonian of the Holy Cross
Mountains and check whether they belong to this genus. First information
about the occurrence of the genus of Aspidichthys in the Holy Cross
Mountains was reported by Gorizdro-Kulczycka (1934) and was later published
by Kulczycki (1957). Kulczycki (1957) described a median dorsal plate and
nuchale plate from the Frasnian deposits of the Kadzielnia Chain (Kielce) and reported the exceptional prominent semidentine tubercles reaching
more than 10 mm in diameter as the most important feature. He also
ascertained that Aspidichthys and Anomalichthys differ
from each other in size and morphology of ornamentation, and these
differences are the result of the North American extent of
Aspidichthys and the European extent of Anomalichthys.
Nevertheless, all specimens formerly belonging of the genus
Anomalichthys are assigned to the genus Aspidichthys
(Denison, 1978).
The stratigraphic occurrence of Aspidichthys is limited to the
Upper Devonian. Most remains have been discovered in the Frasnian, although a few
specimens have been dated as Famennian (Denison, 1978). However, the Holy Cross
Mountains deposits yielding Aspidichthys material are confined to
the Frasnian (Kulczycki, 1957; Narkiewicz et al., 2006).
Institutional abbreviations: WNaZ – Faculty of Earth Sciences, University
of Silesia, Sosnowiec; MWG UW – Faculty of Geology, University of Warsaw;
Muz. PGI-NRI – Geological Museum of the Polish Geological
Institute-National Research Institute, Warsaw.
Anatomical abbreviations: lc – main lateral lines; MD – median dorsal
plate; PDL – posterolateral plate.
Material and methods
The studied material comprises four median dorsal plates of
Aspidichthys ingens Koenen, 1883 and was collected by Jan
Czarnocki in the 1930s, and then described by Kulczycki (1957), Łukasz
Gągała (personal communication, 2001), and Michał Rakociński (personal communication, 2010), as well as the present authors (personal communication, 2015).
Age, locality, and measurements of specimens are listed in Tables 1 and 2.
Specimens are preserved in limestone and prepared at one surface side. In
all cases it is the external side. Specimen Muz. PGI-NRI 5.II.89 was described
and illustrated by Kulczycki (1957, Pl. 11, Fig. 2). Specimen UW MWG
ZI/43/0045 was mentioned by Szrek (2009). Specimens WNaZ/S/4/142 and Muz.
PIG-NRI 1809.II.17 are described for first time. The material is incomplete
but well preserved and reveals diagnostic features like ornamentation and
natural margins of plates. The breadth, length, thickness of plates, height,
and diameter of tubercles were measured (see Table 1). The number of tubercles
on 25 cm2 has been counted as an additional character.
List of the studied specimens and their measurements.
Distribution of the stratigraphical occurrence of described
specimens and diagrams showing Palmatolepis and Icriodus
conodonts percentage share. Abbreviations: E – early; M – middle; L –
late; Pa – Palmatolepis; Icr – Icriodus.
Two limestone samples have been taken from specimens WNaZ/S/4/142 and Muz.
PGI-NRI 5.II.89 and were dated according to conodont assemblage. Each
sample was about 1 kg in weight and was acid-prepared using 10 %
acetic acid. The residuum was separated using a magnetic separator at the
Faculty of Geographical and Geological Sciences of Adam Mickiewicz
University. Conodonts were determined using a
Olympus SZH10 stereoscopic microscope.
Geological setting
The studied area is located in the western part of the Holy Cross Mountains
(central Poland in the Kielce area; Fig. 1). In the studied localities, the Upper
Devonian beds are well exposed. The analysed sections are described by
Szulczewski (1971, Fig. 5), Racki et al. (2002, Fig. 2c), and Szrek and
Ginter (2007, Fig. 7), dated as middle and upper Frasnian based on conodonts
(Table 2).
Location of the investigated outcrops in the Holy Cross Mountains.
(a) Location map of the Holy Cross Mountains (HCM), central Poland (modified
from Kowalczewski, 1971). (b) Diagrammatic cross section through the Holy
Cross Mountains from the Givetian to the top of the Upper Devonian (after
Szulczewski, 1995, modified) with the probable position of the localities
investigated (a – marly limestones and shales; b – condensed cephalopod and
crinoidal limestones; c – marly limestones; d – calcirudites; e – bedded
limestones; f – dolomites; g – massive and bedded limestone; h –
cephalopod limestones; i – clayey and marly shales).
The Late Devonian facies of the area studied is established on the wide
carbonate platform environment, which was divided into blocks during the Frasnian
and stepwise submerged (Szulczewski, 1989, 1995). The submersion of the
carbonate platform was probably an effect of synsedimentary block tectonics
(Narkiewicz et al., 2006; Szulczewski, 1995). The deepening of the Devonian
carbonate platform ended during the Visean and contributed to the
development of a deep-sea pelagic platform (Szulczewski, 1989, 1995;
Szulczewski et al., 1996).
Wietrznia (Fig. 1) is a 30 m deep abandoned quarry located in the
south-eastern part of the city of Kielce, where the Upper Devonian beds are well
exposed. Today the quarry is protected as a geological reserve. The
Aspidichthys remains (Muz. PGI-NRI 5.II.89) were collected in
coarse-grained crinoid limestone and come from the older collection
described by Kulczycki (1957).
The second locality Kowala is a large active quarry situated about 10 km
south of Kielce (Fig. 1). The profile exposed in the quarry comprises the
Frasnian and the Famennian beds representing lithological sets A–L described
by Szulczewski (1971; A–H Frasnian) and Berkowski (2002; H–L Famennian). The
sequence cropping out in the quarry of the Chęciny–Zbrza Basin shows
more than 350 m of the profile (e.g. Racki et al., 2002). The specimens were
collected in marly limestone, which is interbedded with shales (Szulczewski,
1989; Joachimski et al., 2001; Racki et al., 2002).
Specimens from Wietrznia and Kowala were found in debris, and conodont
analysis was necessary to determine age; however, it was possible to determine
their position precisely in the profile. The limestone from Wietrznia quarry
was dated as Palmatolepis punctata (middle Frasnian) and was
described by Szulczewski (1971, Fig. 5). The conodont zone analysis of marly
limestone from Kowala quarry showed Palmatolepis rhenana zone
(upper Frasnian) and was described by Racki et al. (2002, Fig. 2c).
The last locality is Płucki near Łagów, about 36 km east of Kielce (Fig. 1).
There are two outcrops located in the south-eastern neighbourhood of the village of Łagów, on the slope of Łagowica Valley, in the eastern fragment
of the Kielce–Łagów Synclinorium. In the Upper Devonian sequence, the
so-called Łagów Beds consist of carbonate rocks (limestones and
shales) as well as distinctive horizons of lower and upper Kellwasserkalk built up
with classic black bituminous limestones. These two horizons are thought to
be connected with anoxic episodes during the Late Devonian (Janiszewska et al.,
2007; Szrek, 2007; Szrek and Ginter, 2007, 2008; Rakociński et al., 2016).
Fish remains from Płucki are best preserved within the collection of all
Aspidichthys specimens and were collected in situ from the
section. They occur in the outcrop with the upper Kellwasserkalk horizon.
The outcrop is about 3 m deep and exposes limestones, marly limestones,
marls, and shales. The sequence is strongly disturbed by the presence of a
Devonian underwater landslide ascertained very recently
(P. Szrek and S. Salwa, personal communication, 2014). The black Kellwasserkalk
limestone is about 0.5 m thick and consists of several discontinuity events
(Janiszewska et al., 2007; Szrek, 2007; Szrek and Ginter, 2007, 2008;
Rakociński et al., 2016). The lower, entirely Frasnian part (up to
30 cm) is characterized by a marl at the bottom with gradual changes
upwards into limestone. Fish fossils are quite common, and large armours of
Aspidichthys placoderms are most numerous. The middle part (up to 10 cm)
is the most fossiliferous and is particularly rich in cephalopods and minute
arthrodire placoderms. According to precise analysis of Janiszewska et al. (2007)
the Frasnian–Famennian boundary is situated within this part of the
horizon. The uppermost part entirely belongs to the Famennian and is
characterized by numerous invertebrates and fishes, but no
Aspidichthys was ascertained here.
Type species: Aspidichthys ingens von Koenen, 1883Aspidichthys ingens von Koenen, 1883(Figs. 2–5)1933 Anomalichthys magnus n. sp., W. Gross, pp. 42–44, Tab. 6, Fig.
4.1933 Aspidichthys? ingens. – W. Gross, pp. 48–49, Tab. 6, Fig. 5.1937 Aspidichthys ingens. – W. Gross, p. 40, Pl. 5, Fig.
4.1956 Aspidichthys clavatus. – G. W. Sinclair & D. R.
Walker, pp. 135–137, Figs. 1–3.1957 Anomalichthys ingens. – J. Kulczycki, pp. 333–335, Figs. 1–3.1973 Aspidichthys cf. ingens Koenen, H.-P. Schultze, pp.
69–75, Pl. 2A–D.1976 Aspidichthys. – J. P. Lehman, pp. 21–22, Pl. 9, Figs. C–D.1978 Aspidichthys ingens. – R. Denison, p. 103, Fig. 84.Material: Four fragments of median dorsal plates: Muz. PGI-NRI 5.II.89 (Kulczycki, 1957,
Fig. 2), MWG UW ZI/43/0045 (Figs. 2–3), Muz. PIG 1809.II.17 (Fig. 4), and an
incomplete median dorsal plate WNaZ/S/4/142 (Fig. 5). See Table 1 for
detailed measurements and Table 2 for horizons.Locality and age: Muz. PGI-NRI 5.II.89: middle Frasnian (Palmatolepis punctata
conodont zone) of Wietrznia Quarry in Kielce; WNaZ/S/4/142: upper Frasnian
(Palmatolepis rhenana conodont zone) of Kowala Quarry; Muz. PIG
1809.II.17, MWG UW ZI/43/0045: upper Frasnian (Palmatolepis linguiformis conodont zone) of Płucki near Łagów.Description: Specimen Muz. PGI-NRI 5.II.89 (Kulczycki, 1957, Pl. 9, Fig. 2) shows the
anterior part of a median dorsal plate with clearly visible ornamentation. The
median dorsal is broken and lacks its natural boundaries; the carinal process
on the visceral side typical of brachythoracids is not visible. The
ornamentation consists of smooth and rounded tubercles that are irregularly arranged.
The diameter of tubercles varies between 2 and 4 mm, and the height varies
between 1.8 and 3 mm. The number of tubercles on 25 cm2 of the shield
is approximately 101.
Specimen no. MWG UW ZI/43/0045. (a) Dorsal view of the specimen
(arrow shows anterior part), (b) lateral view of the reconstructed armour
(preserved parts are marked), and (c) dorsal view of specimen with particular
plates and elements marked with lines.
Specimen no. MWG UW ZI/43/0045. Magnified anterior part of the
median dorsal plate in (a) lateral and (b) dorsal view.
Specimen no. Muz. PIG 1809.II.17. (a) Dorsal view of the median
dorsal plate (arrow shows anterior part), (b) median dorsal plate in anterior
view, (c) one of the armour plates (probably the posterior lateral
plate), and
(d) magnified area of the median dorsal plate showing the ornamentation.
Specimen no WNaZ/S/4/142, anterior part of the median dorsal
plate. (a) Dorsal view, (b) lateral view (arrow shows anterior part), (c) anterior
view, and (d) magnified area of the median dorsal plate showing the ornamentation.
This specimen described above was taken into consideration by Kulczycki
(1957) but not illustrated. Nevertheless, it served as the source of
material from which conodonts were prepared. Specimens described below were
collected very recently and have not been published before.
The specimen MWG UW ZI/43/0045 (Figs. 2–3) shows partially preserved median
dorsal plate and left posterior dorsolateral plate. The plate is strongly
arched, enclosing at an angle of ca. 120∘ in the anterior half and
rounded and flattened out in the posterior half. The external side of the plate
is covered with large and rounded tubercles irregularly arranged. Tubercles
are 5 mm in diameter at their largest and 1.4 mm in height. The number of
tubercles on 25 cm2 was not counted because a large part of them are
eroded. The left posterior dorsolateral plate is preserved in the same
block of rock and could belong to the same individual. This plate has the
natural boundary along the lower, left, and upper margin, the last of which being an
overlapping surface for the median dorsal plate (Fig. 2). The lateral line
is well preserved, especially on left half of the plate, and is visible from
the left anterior corner, close to the upper margin and border with median
dorsal plate for about one-third of its length, and then turns and extends onto
the right margin in approximately the middle of the plate. The posterior
part of this plate is still covered by the matrix, and the most posterior margin
lies under the anterior part of the median dorsal plate. The course of the
lateral line is additionally marked with a longitudinal arrangement of
tubercles.
Specimen no. Muz. PGI-NRI 1809.II.17 (Fig. 4) is represented by a median
dorsal plate. The plate is very fissured and curved. The angle between the
left and right sides is 120∘ in the anterior half (Fig. 4b); the
posterior part is flattened secondarily. On the lateral part of the shield a
natural boundary is preserved, but the keel is invisible. The shield is
ornamented with irregularly arranged and not widely removed from each of the other
tubercles (Fig. 4d). The tubercles are rounded and are 2 to 2.5 mm in diameter. They
are also more or less the same sizes on the entire surface of the shield.
The height of tubercles was not measured because of the high abrasion of
them. The number of tubercles on 25 m2 of the shield is 72. The
specimen additionally included a fragment of unidentified other plate,
probably a part of one of the lateral plates of the trunk shield (breadth
3.65 cm,
length 5.5 cm) with the same type of ornamentation. In the lateral and
posterior regions of the median dorsal plate they are larger and sometimes
also irregularly confluent.
The specimen (WNaZ/S/4/142, Fig. 5) shows the anterior part of the median dorsal
plate. The plate is thick (about 4.7 cm) and the visceral surface is not
visible. The natural anterior margin is preserved. The ornamentation
consists of round and smooth tubercles without regular arrangement.
Tubercles are about 1 to 3 mm wide in diameter and 1.5 cm in height. The
number of tubercles on 25 cm2 have not been counted because a large
part of them are eroded.
In the anterior part of those plates (MWG UW ZI/43/0045, Figs. 2–3; WNaZ/S/4/142, Fig. 5), the small crest occurs. The crest is in the form of a
median longitudinal thickening of steep sides, ascending gradually and
reaching a height of about 25 mm in the anteriormost margin. The symmetry plane of
this crest meets the plane of the shield at an angle of 30∘. The
surface of this crest is covered with tubercles similar to the rest of the
shield. The crest was originally present on specimen Muz. PGI-NRI
1809.II.17 but has been lost because of damage to this region of the shield (Fig. 4a).
Counting of tubercles on the described specimens revealed that a younger specimen
from Płucki (Muz. PGI-NRI 1809.II.17) has a lower number of tubercles than
the older specimen from Wietrznia Quarry (Muz. PGI-NRI 5.II.89) (see
Table 1).Discussion: Already known species of the genus Aspidichthys are represented by
A. ingens Koenen, 1883 and A. clavatus Newberry, 1873 (see
the discussion by Denison, 1978, p. 103). The presented material is similar in
shape and morphology to the specimens described by Kulczycki (1957) but is
characterized by smaller tubercles. Tubercles in Kulczycki's (1957) material
reach up to 10 mm, while in the studied material they do not extend to even 5 mm in diameter.
On specimen WNaZ/S/4/142, tubercles reach the smallest sizes even though this
specimen represents the largest individual among all analysed specimens.
The characters observed on studied specimens are very peculiar and correspond well to randomly distributed tubercules. Tubercules have stellate base of 5 mm in diameter at their largest. The plate is thick, arched and elongated. Those features have been observed and illustrated by previous authors (Newberry, 1873,
Pl. 35, Fig. 1; Whiteaves, 1892, p. 355; Gross, 1933, Tab. 6, Fig. 4; Sinclair
and Walker, 1956, Fig. 1–3; Kulczycki, 1957, Pl. 1–3; Schultze, 1973), as well as
preliminary characterized material from the Holy Cross Mountains (Szrek,
2009). All specimens described herein are different from those discussed by
Schultze (1973, Pl. 2, Fig. a–b) by clearly larger distances between
tubercles and thus are more reminiscent of those illustrated by Newberry
(1873, Pl. 35, Fig. 1), Gross (1933, Tab. 6, Figs. 4–5), Sinclair and Walker
(1956, Fig. 1), and Kulczycki (1957, Pl. 11, Figs. 2–3). Differences could
lie in a local palaeogeographic morphological variations suggested by
Kulczycki (1957) rather than sizes of particular individual and species
differentiation.
The specimens where the anterior margin of the median dorsal plate is
preserved (MWG UW ZI/43/0045, Figs. 2–3; WNaZ/S/4/142, Fig. 5a–c) show a
more rounded margin in the part where it turns posteriorly than on the
specimen illustrated by Sinclair and Walker (1956, Figs. 1–2), on which this
part turns at almost a right angle. This region is visible only in the
specimens MWG UW ZI/43/0045 and WNaZ/S/4/142 and those of Sinclair and Walker
(1956, Figs. 1–2). Moreover, the carinal process mentioned by Kulczycki
(1957) and Schultze (1973) is not accessible for investigation because it is not
visible in the new material from the Holy Cross Mountains, making a detailed
comparison difficult. However, other morphological features of the external
side of the median dorsal plates (overall shape longer than wide,
ornamentation composed of large tubercles, thickness of bones that
are arched and which enclose at an angle of 120∘ of the median dorsal
plate) correspond to other described specimens (Sinclair and Walker,
1956; Kulczycki, 1957) and confirm identification of Aspidichthysingens Koenen, 1895.
The median dorsal plate of species Aspidichthys ingens represented
by specimen no. WNaZ/S/4/142 is not complete, but species-related features
are preserved (e.g. large, rounded irregularly arranged tubercles and
relatively thick shield; Denison, 1978) which are commonly observed on the
plates and suggest that the plate belongs to this species. The studied
specimen has smaller tubercles (1–3 mm in diameter) than the specimens
described by Kulczycki (1957), which have tubercles of even 10 mm in diameter.
This specimen can be compared with Aspidichthys described by
Schultze (1978, Pl. 3), where the ornamentation contains tubercles about 4 mm
in diameter; however, the studied specimen shows a large keel. In the
material described by Schultze (1978), an inconsiderable bump occurs on the
anterior part of the shield. This bump is clearly visible on almost all specimens
studied.
The studied remains occur in the pelagic facies, suggesting that
Aspidichthys lived in a deep-water environment (Ivanov and Ginter,
1997; Szrek, 2006) as active predators would have had a demersal lifestyle
(Janiszewska et al., 2007; Szrek, 2007). The conodont analysis shows that
placoderms from Wietrznia Quarry lived in slightly shallower water than
those from Kowala Quarry. In the sample from the
punctata zone (Wietrznia Quarry), Icriodus is more numerous
than in the sample from the early rhenana zone (Kowala Quarry), where
Icriodus elements are in the minority, which suggests a deeper
environment. In the sample from Wietrznia Quarry (punctata
conodont zone) there were 10 Icriodus elements and 19
Palmatolepis elements. In the sample from Kowala Quarry
(rhenana conodont zone) there were 25 elements of Icriodus and
326 elements of Palmatolepis. Sample form Płucki revealed 33
Icriodus and 44 Palmatolepis elements. For details, one
can refer to Table 2, which indicates a progressive and stepwise deepening
of the environment and is correlated with the synsedimentary block tectonics
and permanent growth of the sea level in the studied area during the Frasnian
suggested by previous authors (e.g. Szulczewski, 1995). Analyses of the Płucki
section (Racki et al., 2002; Woroncowa-Marcinowska, 2006; see Table 2)
demonstrated a deep-sea environment in the linguiformis conodont zone.
Moreover, the regression at the Frasnian–Famennian boundary has been
suggested based on the percentage share of Palmatolepis and
Icriodus elements (Janiszewska et al., 2007), which could be
correlated with global changes in sea level (McGhee Jr., 2013). The
conjunction of the regression and environmental changes during the reef fauna
crisis could have led to the disappearance of Aspidichthys during
the Frasnian to Famennian transition.
The analysis of the placoderm fauna from the Upper Kellwasserkalk of Płucki shows similarities to the contemporary fauna from Bad Wildungen. For 14
genera ascertained in Płucki (Szrek, 2008, 2009), 9, including
Aspidichthys, are common to Bad Wildungen. This is most likely
directly related to the short palaeogeographic distance between both
localities. Exhaustive analysis of palaeobiogeographic distribution of
different placoderm taxa also from Morocco, France, Iran, and Australia
suggests vicinities between all of these areas without a significant barrier
(Rücklin, 2010) for bottom-dwelling fishes (e.g. deep ocean).
Conclusions
Detailed description of Aspidichthys placoderm from the
Kellwasserkalk locality in the Holy Cross Mountains is provided for the
first time. The remains constitute the most important placoderm assemblage
associated with a single genus from the Devonian of the Holy Cross Mountains.
The described specimens belong to Aspidichthys ingens Koenen, 1883.
Previous stratigraphic occurrences of specimens mentioned by
Gorizdro-Kulczycka (1934), Kulczycki (1957, p. 287), and Ivanov and Ginter
(1997, Table 1) based on old lithostratigraphic zonation are now updated to
standard conodont zonation (Ziegler and Sandberg, 1990). Analysis shows that
stratigraphic occurrence of this genus is limited to the Frasnian, in the Holy
Cross Mountains.
Environmental analysis based on conodont biofacies shows that
Aspidichthys placoderms have adapted to a deepening environment
as their fossils occur in Icriodus-dominated as well as in
Palmatolepis-dominated facies. The occurrence of the same taxa in
many different locations (Bad Wildungen, Morocco, North America, Iran, Holy
Cross Mountains) is in agreement with the absence of a biogeographic barrier
effectively separating particular ecosystems postulated by Rücklin
(2010).
Data availability
Specimens described herein are housed at the following institutions:
Geological Museum of the Polish Geological Institute-National Research Institute in Warsaw, Poland (collection number Muz. PGI-NRI 5.II and 1809.II);
Faculty of Geology, University of Warsaw, Poland (collection number MWG UW
ZI/43); and Faculty of Earth Sciences, University of Silesia in Sosnowiec, Poland (collection number
WNaZ/S/4).
Acknowledgements
We thank Vincent Dupret and the anonymous reviewer for very useful
revision and helpful comments. We are grateful to M. Rakociński
(University of Silesia, Sosnowiec) and Ł. Gągała (Georex,
Argenteuil) for providing a part of studied collection; to Z. Bełka (Adam
Mickiewicz University, Poznań) for help during conodonts identification
and to B. Berkowski (Adam Mickiewicz University, Poznań); and G. Gierliński
(Polish Geological Institute-National Research Institute,
Warsaw), who kindly revised an early version of the manuscript. Piotr Szrek was funded by
a Polish Geological Institute – National Research Institute (grant number
61.2401.1301.00.0).
Edited by: F. Witzmann
Reviewed by: V. Dupret and one anonymous referee
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