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An International Journal of Paleobiology
ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20
Fishes from the Miocene lacustrine sequence of
Oleksandr Kovalchuk, Adam Nadachowski, Ewa Świdnicka & Krzysztof
To cite this article: Oleksandr Kovalchuk, Adam Nadachowski, Ewa Świdnicka & Krzysztof
Stefaniak (2019): Fishes from the Miocene lacustrine sequence of Bełchatów (Poland), Historical
To link to this article: https://doi.org/10.1080/08912963.2018.1561671
Published online: 09 Jan 2019.
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Fishes from the Miocene lacustrine sequence of Bełchatów (Poland)
, Adam Nadachowski
, Ewa Świdnicka
and Krzysztof Stefaniak
Department of Aquaculture, National University of Life and Environmental Sciences of Ukraine, Kyiv, Ukraine;
Department of Palaeontology,
National Museum of Natural History of the National Academy of Sciences of Ukraine, Kyiv, Ukraine;
Institute of Systematics and Evolution of
Animals of the Polish Academy of Sciences, Kraków, Poland;
Department of Palaeozoology, Institute of Environmental Biology, University of
Wrocław, Wrocław, Poland
The paper deals with results of the study of freshwater ﬁsh remains obtained from the Miocene deposits
of Bełchatów open-cast brown coal mine in 1984 and 1986. These fossils are represented mainly by
numerous disarticulated bones which are assumed as belonging to cyprinid (Tinca sp.) and esocids
(Esox sibiricus, Esox sp.), as well as to indeterminate bony ﬁshes (Teleostei indet.). The ﬁnd of the
Siberian pike described herein is the oldest occurrence of this species in Europe, and allows extending
its previously known range to the west. Some aspects of the palaeogeography and palaeoecology of
the late early Miocene ﬁsh assemblage of Bełchatów with reference to the distribution of E. sibiricus are
Received 6 July 2018
Accepted 18 December 2018
dynamics; brown coal mine;
The composition of the freshwater ichthyofauna of the
Northern Hemisphere was formed in general terms during
the late Cenozoic (Sytchevskaya 1989). Miocene was a period
when the seasonality of climate has increased, however, the
average annual temperature has been relatively stable
(Herbert et al. 2016). It was marked by profound environ-
mental change in ecosystems, in particular the appearance of
the latitudinal gradient visible in modern times (Ivanov et al.
2011). The faunistic analysis of the fossil ﬁsh remains from
the heterochronous deposits in Central Europe allows us to
trace the temporal changes in ﬁsh assemblages, to clarify the
time of the appearance of extant taxa and their groups in the
fossil record, to ﬁnd out the routes of their migrations, to
indicate the centres of origin of individual components of the
assemblages and the formation of their ranges.
In this paper, we provide a detailed description of all
freshwater ﬁsh fossils obtained from the Miocene deposits
of the Bełchatów lignite mine in Poland. This locality was
discovered in the 1960s, and later, since 1980s, numerous
fossils, namely plants (Worobiec 1995; Worobiec and
Worobiec 2005; Worobiec and Szynkiewicz 2007), mollusks
(Stworzewicz 1995,1999a,1999b; Stworzewicz and Sołtys
1996; Kadolsky and Piechocki 2000), mammals (Kowalski
1993a,1993b,1997; Rzebik-Kowalska 1993,1994,1996,
2005; Nadachowski 2001; Garapich 2002) were found there
and described in a huge number of publications (for other
references, see also Fostowicz-Frelik et al. 2012).
We attempt to clarify the systematic position for ﬁsh
remains brieﬂy described by Jerzmańska and Hałuszczak
(1986) as well as those not processed before. All studied
specimens come from several exposures located on the
exploited escarpment in the Bełchatów mine. Based on
obtained data, we provide a short overview of the environ-
ment of Bełchatów, with an insight to its palaeobiogeography.
The studied locality is situated in central Poland (51°15ʹN,
19°20ʹE), between Warta and Pilica rivers, near 15 km
southwards of the town Bełchatów (Figure 1(A)). This is
an open-cast brown coal mine, whose deposits are geneti-
cally connected with a series of tectonic depressions within
the Kleszczów Graben (Stuchlik et al. 1990;Krzyszkowski
1993; Szynkiewicz 2000;GotowałaandHałuszczak 2002).
That graben is ﬁlled with Neogene and Quaternary deposits
lying on Jurassic and Cretaceous rocks, and a salt diapir of
Permian (Zechstein) age in its central part (Hałuszczak
2004). The Neogene sequence in Bełchatów consists from
bottom to top of subcoal (PW), coal (W), clayey-coal (I-W),
and clayey-sandy (I-P) lithostratigraphic units (Figure 1(C)).
The formation age of the entire thickness of clastic deposits
intercalated with lignite beds, lake marls and volcanic ash
layers near 60 m in depth is estimated to be early to early
late Miocene, i.e. Eggenburgian–Pannonianinthemarine
scheme of Central Paratethys stages (Fostowicz-Frelik et al.
2012). Three principal faunal levels have been previously
recognised (Nadachowski 2001). Among them, the oldest is
Bełchatów C or BE-C (Ottnangian–Karpatian, 18.1 ± 1.7 Ma
after Burchart et al. 1988; Piller et al. 2007;Hilgenetal.
(early Badenian, 16.5 ± 1.3 Ma according to Burchart et al.
1988; Piller et al. 2007;Hilgenetal.2012), and the most
recent is those of Bełchatów A (BE-A). Based on small
mammalian remains, the age for the latter assemblage can
be correlated to late Sarmatian–early Pannonian (Kowalski
CONTACT Oleksandr Kovalchuk firstname.lastname@example.org
© 2018 Informa UK Limited, trading as Taylor & Francis Group
Published online 09 Jan 2019
and Rzebik-Kowalska 2002; Fostowicz-Frelik et al. 2012).
Fish-bearing strata are located just between the BE-C and
BE-B assemblages, being younger than BE-C and older than
BE-B.Theageoftheﬁsh fossils is late early Miocene (see
Material and methods
The fossil ﬁsh material from the beds of Bełchatów was
obtained by A. Jerzmańska, E. Świdnicka, accompanying
with A. Hałuszczak and R.J. Wilczyński (both geologists)
during their ﬁeld campaigns in 1984 and 1986 (Figure 1(B)).
It comes from exposures near boreholes Nos. 88/14, 87/13
and 341B and was collected at +50 and +62 meters above sea
level (a.s.l.). The ﬁsh fauna of Bełchatów occurs in the higher
part of the coal unit (W) consisting of three coal sems (i.e.
main coal (D) and C (III) and B (II)), two levels of humic
coals (called as ‘cuboidal clays’), lacustrine limestones and
clayey layers (Figure 1C).
The environmental setting of these layers was previously
interpreted as a shallow lake with a thick layer of organic
debris accumulated at the bottom (Wagner 1996; Widera
2016). The burial of remains most likely occurred in anoxic
conditions (Jerzmańska and Hałuszczak 1986). In total, 140
disarticulated bones and their fragments embedded in the
coaly clay matrix from Bełchatów are stored in the
Department of Palaeozoology (Institute of Environmental
Biology, University of Wrocław). Bones and scales were
identiﬁed based on comparisons with extinct and modern
taxa from the literature (Galkin 1958; Obrhelová 1970;
Sytchevskaya 1974,1976,1980,1989; Gaudant 1978;
Gaudant et al. 2002; Kovalchuk et al. 2017) and osteological
database (Tarcerie et al. 2016). Determination of elements
was accomplished using diagnostic features based on com-
Figure 1. Geographic location and stratigraphical position of the Bełchatów Lignite Mine (after Jerzmańska and Hałuszczak 1986, with modiﬁcations): A–location of
the Bełchatów in Poland; B–collecting of ﬁsh remains in Bełchatów by A. Jerzmańska and A. Hałuszczak in 1984; C–generalized scheme of the proﬁle. For
abbreviations –see Geological setting.
2O. KOVALCHUK ET AL.
parative observations and literature summaries (Grande
1999; Grande et al. 2004). There are numerous additional
small bone fragments in the palaeoichthyological collection
of ZPALWr. (see Appendix). However, due to their poor
preservation, these specimens have been neither identiﬁed at
least to order or family, nor adequately measured. The
taxonomical hierarchy in this paper follows Nelson et al.
(2016), Froese and Pauly (2018). The osteological terminol-
ogy in this paper follows Sytchevskaya (1976) and Lepiksaar
(1994). Bones were measured according to Morales and
Rosenlund (1979) using an electronic caliper Mitutoyo with
0.01 mm precision.
Abbreviations:BM–acronym of ichthyological collection
from the Miocene deposits of Bełchatów; ZPALWr. –
Department of Palaeozoology, University of Wrocław.
Subdivision Teleostei Müller 1846 sensu Arratia 1999
Order Esociformes Berg 1940
Family Esocidae Cuvier 1817
Genus Esox Linnaeus 1758
†Esox sibiricus Sytchevskaya 1974
Figure 2(A–K, M)
Esox sp.: Jerzmańska and Hałuszczak 1986, p. 26, Figure 2.
Two vomers (ZPALWr. BM/59, BM/77), one frontal (ZPALWr.
BM/89), three parasphenoid fragments (ZPALWr. BM/39, BM/
67-68), one left palatine in part (ZPALWr. BM/15) and counter-
part (ZPALWr. BM/16), four fragments of palatine (ZPALWr.
BM/43-46), one quadrate (ZPALWr. BM/76), three articulars
(ZPALWr. BM/22-23, BM/114), one left dentary (ZPALWr.
BM/10), 12 dentary fragments (ZPALWr. BM/11-12, BM/17,
BM/19-20, BM/26-27, BM/38, BM/71-74), one isolated tooth
(ZPALWr. BM/13) and its fragment (ZPALWr. BM/79), two
fragments of cleithra (ZPALWr. BM/78, BM/101).
The vomers (Figure 2(A,B)) are represented by two notably
massive vomerine heads, each with partly or completely bro-
ken arm. The latter is supposed to be long, wide and ﬂattened,
as far as can be judged basing on morphology of this bone in
other fossil and modern pikes. Well-developed rounded pro-
jections are located anterolaterally on edges of the dental ﬁeld
whose width in BM/59 and BM/77 is 18.2 mm and 17.0 mm,
respectively. Teeth are bigger in the anterior part of the bone
as compared with those located posteriorly.
The frontal (Figure 2(C)) is characterized by a thickened
posterior portion of the medial suture, a narrow sphenotic slit
and a distinct anterolateral crest extending from the shaft of
the supraorbital canal on the ventral surface. Lateral edges of
the bone are slightly eroded.
Three parasphenoids (Figure 2(D)) are preserved in part,
with clearly visible grooves of the carotid arteries. Posterior part
of the bone is widened caudally, while its dorsal medial crest is
narrowed in the same direction. There is a medial denticle at
the level of ascending wings. Width of the bone in its medial
part varies from 5.5 mm in BM/39 to 12.4 mm in BM/67.
The palatine (Figure 2(K)) is wide and ﬂattened, with
destroyed edges. There are seven rows of C-shaped small
tooth bases on the ventral side of the palatine. Width of the
bone is 12.7 mm.
The quadrate (Figure 2(M)) is destroyed and characterized
by the presence of wide articular facet.
The articular bone (Figure 2(I)) is massive, with a high wall
(anguloarticular angle is equal to 70°), an elongated and later-
ally compressed posterior process, and a wide articular facet.
The dentary (Figure 2(E,F, J)) is elongate and massive. It
has a well-developed dental shelf, which smoothly rises onto
the medial wall. Teeth at the symphysis (Figure 2(J)) are
arranged in two rows, and internal one is better developed.
Twenty C-shaped crown bases, gradually increasing in dia-
meter in the direction towards the symphysis, are well visible
at the bone surface. A clear lateral groove separates the ante-
rior edge of the dentary from rest of the bone and forms
a ventral bulge in the symphysis. The ﬂattened anterior edge
of the bone is almost straight and thickened at the symphysis.
The latter is characterized by the absence of the symphysial
notch. Six pores for the sensory canal (Nelson 1972) are
visible at the ventral surface of the dentary. Measurements
of specimens from Bełchatów are presented in Table 1.
There is one elongate, dagger-shaped marginal tooth
(Figure 2(G,H)). It is laterally compressed, with slightly
curved crown, partly worn sharp tip, and rounded
(C-shaped) inﬂated base. Cutting edges are slightly narrowed.
Height of the tooth is more than 12 mm, width –6.8 mm.
Remains described above are identical in overall morphol-
ogy with those of the extinct Siberian pike Esox sibiricus
previously known in the fossil record of Kazakhstan
(Sytchevskaya 1976), Mongolia (Sytchevskaya 1989)and
Ukraine (Kovalchuk et al. 2017). Concerning the measure-
ments, bones from Bełchatów are signiﬁcantly larger than
other fossils previously assigning to this species. It diﬀers
from modern Eurasian pikes and American pickerels in
shortened, wide and ﬂattened palatine, double tooth row at
the symphysis, more massive dentary, larger anguloarticular
angle, as well as wider facet of the quadrate. Siberian pike
resembles the Amur pike E. reichertii (and opposed to
E. lucius) in morphology of the vomer and frontals, orienta-
tion of the maxillary process of palatine, and in the structure
of the anterior median crest of the parasphenoid. At the
same time, Siberian pike diﬀers from E. reichertii in more
robust, shorter and higher dentary with straight symphysial
and wider palatine, as well as higher anguloarticular angle.
As for the extinct species, E. sibiricus diﬀers from
E. aralensis in narrower dental shelf, absence of the sym-
physial notch and two-row dentition (Sytchevskaya 1976).
Siberian pike resembles E. moldavicus in the structure of the
dentigerous surface of the palatine, but diﬀers in the pre-
dominance of the two-row dentition at the dentary symphy-
sis. It is not possible to adequately compare our material
HISTORICAL BIOLOGY 3
with E. aralensis duetotheabsenceofrespectiveskeletal
elements in Bełchatów.
One vomer fragment (ZPALWr. BM/102), ﬁve dentary fragments
(ZPALWr. BM/24-25, BM/28, BM/32, BM/112), eight tooth
fragments (ZPALWr. BM/47, BM/60, BM/75, BM/107-111),
one opercular fragment (ZPALWr. BM/21), two subopercles
(ZPALWr. BM/33, BM/106), 14 abdominal centra (ZPALWr.
BM/14, BM/34-37, BM/40-42, BM/84-88, BM/104), one sample
with scale plates (ZPALWr. BM/18).
Vomers, articular, opercular and subopercular bones
(Figure 2(L)), dentaries and cleithra contain some features
Figure 2. Miocene ﬁsh remains from Bełchatów. A-K, M –Esox sibiricus Sytchevskaya (1974): A–vomer ZPALWr. BM/77, toothed surface; B–vomer ZPALWr. BM/59,
toothed surface; C–frontal ZPALWr. BM/89, in medial view; D–parasphenoid fragment ZPALWr. BM/68, in dorsal view; E-F –dentary ZPALWr. BM/10, in ventral (E)
and dorsal (F) view; G-H –isolated tooth ZPALWr. BM/13, in lateral (G) and posterior (H) view; I–articular fragment ZPALWr. BM/114, in medial view; J–dentary
ZPALWr. BM/38, in dorsal view; K–palatine ZPALWr. BM/15, toothed surface; M–quadrate ZPALWr. BM/76, in lateral view. L, N-Q –Esox sp.: L–subopercle
ZPALWr. BM/33, in lateral view; M–dentary fragment ZPALWr. BM/112, in lateral view; O–sample with scale plate ZPALWr. BM/18; P-Q –abdominal centrum
ZPALWr. BM/40, in anterior (P) and lateral (Q) view; R-S –Tinca sp., opercular fragment ZPALWr. BM/100, in medial (R) and lateral (S) view. Scale bar equals 5 mm in
B-C, G-H, L, N-O, 1 cm in A, D, I-K, M, P-S, and 2 cm in E-F.
4O. KOVALCHUK ET AL.
characteristic of Esox (see Codrea et al. 2018 for more
details). The teeth at the symphysis in one dentary
ZPALWr. BM/112 (Figure 2(N)) are arranged in one row
whose width is equal to 2.2 mm. Isolated teeth are repre-
sented by their medial portions. Centra (Figure 2(P,Q)) are
circular in end view. They are characterized by deep acces-
sory pits on either side of the central longitudinal bony
strut on the lateral surface. The latter is composed of
longitudinal bony ﬁbres. Diameter of preserved centra var-
ies from 13.0 to 24.9 mm (mean –18.8 mm). Scales (Figure
2(O)) are typical to those in pikes. Their plates are elon-
gated oval in shape, with strongly developed anterior ﬁeld
bearing three rounded festoons. The core of these scales is
sharply shifted to the posterior edge.
The specimens share similar morphology with respective
bones in Esox. However, their species attribution is uncertain
due to the lack of reliable diagnostic characters.
Order Cypriniformes Bleeker 1859
Family Cyprinidae Bonaparte 1832
Genus Tinca Cuvier 1817
Cyprinidae gen. indet.: Jerzmańska and Hałuszczak 1986,p.26.
One opercular fragment (ZPALWr. BM/100).
The opercle is represented by its half part, with the wide
concave lamina. It is supposed that the bone has been trape-
zoid in shape. Anterior margin of the opercle is almost
straight, the dorsal one is slightly convex, while the posterior
and ventral margins are completely broken. The supraglenoi-
dal process with widened base (~2.6 mm) is well pronounced,
and the articular facet is lensed in shape and narrowing at the
top. Width of the facet is 3.9 mm, and depth is near 5.2 mm.
In lateral view, a distinct narrow groove runs parallel to the
anterior margin of the bone. Assuming the proportions of the
articular facet and the total dimensions are the same as in
Recent tench Tinca tinca, reconstructed length of the opercle
is about 58 mm, while its width is 42 mm.
The opercle resembles those of Tinca in the shape of its anterior
margin and articular facet (see, for example, Radu 2005,p.57),
as well as in the structure of internal postarticular crests and
supraglenoidal process (Obrhelová 1970,p.143,ﬁgure 25A, B).
However, we identiﬁed the specimen as belonging to this genus
only tentatively because of its poor preservation (i.e. destroyed
both posterior and ventral margins).
The fossil fauna of Bełchatów contains at least three ﬁsh taxa.
Two of them (Esox sibiricus, Tinca sp.) are clearly distin-
guished. The Siberian pike have presumably dominated in
this assemblage. Presumably, the poor diversity of the fossil
assemblage reﬂects the poor diversity of the ﬁshes. Such
a small species diversity allows us to assume that these ﬁshes
lived in a stressed environment as suggested by Kadolsky and
Piechocki (2000) for aquatic snails.
It is important to focus on some ecotopic preferences of the
extant representatives of the identiﬁed genera. Pike is an
ectotherm ﬁsh with a wide range of environmental tolerances
(Casselman and Lewis 1996; Newbrey et al. 2008). It is character-
ized as a keystone piscivore shaping the composition, abundance
and distribution of ﬁsh assemblages (Craig 2008). This ﬁsh most
commonly occurs in shallow, moderately productive and vege-
tated waters (Diana 1979; Harvey 2009). Tench is an omnivorous
ﬁsh with a broad diet. It tends to feed in areas with a large supply
of macrophytes (Nordstrom 2011). This ﬁsh prefers standing to
slowly ﬂowing waters and muddy bottom with abundant vegeta-
tion (Moreno Rendón et al. 2003). According to Worobiec (1995,
2003),theﬂora of Bełchatów is represented by plant remains of 21
species belonging to 15 genera of gymnosperms and angiosperms.
Most of them are representatives of the Arcto-Tertiary subtropical
ﬂora (Worobiec 1995,2003). The ﬁnding of ﬁsh fossils in humic-
coal clayey deposits also suggests shallow lake conditions
(Worobiec ; Widera 2016).
Diversity of cyprinids and esocids in the Miocene strata of
Europe sheds light upon the evolutionary history of these ﬁshes
and establishing some characteristics of their distribution dur-
ing the late Cenozoic (Sytchevskaya 1989). The genus Tinca is
well-known in the European Neogene fossil record (Kovalchuk
2015). Tench remains described are known from the
Burdigalian of Czech Republic (Weiler 1966)andFrance
(Ginsburg et al. 2000), Serravallian of Germany (Böhme;
Gaudant et al. 2002), as well as from the Tortonian and
Messinian deposits of Austria, Moldova, Russia, Slovakia
(Sytchevskaya 1989;Gaudant1994; Böhme and Ilg 2003;
Titov et al. 2006; Kovalchuk et al. 2014a,2014b) and Ukraine
(Kovalchuk 2015,2017). The discovery of numerous remains
assigned to E. sibiricus in the fossil record of Bełchatów has
a considerable biogeographical signiﬁcance (Figure 3).
Table 1. Measurements of dentaries of †Esox sibiricus Sytchevskaya (1974) from
Bełchatów, in mm.
of dentary near the
Width of tooth
BM/11 16.3 10.9 3.5
BM/12 –14.7 5.0
BM/38 11.4 7.3 4.5
HISTORICAL BIOLOGY 5
This extinct pike was previously reported from the
Serravallian beds of Kazakhstan, as well as from the
Tortonian and Messinian of Mongolia, Russia and Ukraine
(Sytchevskaya 1976,1980,1989; Kovalchuk et al. 2017).
Therefore, Bełchatów, being equal in age to those from the
early Miocene of West Siberia (Sytchevskaya 1976), is the
oldest occurrence of Esox sibiricus in Europe, and now it is
the most western ﬁnd for this species. We assume that the
range of the Siberian pike during Miocene covered a wide
zone from Central Asia to Central Europe, between 40° and
60° of northern latitude (Figure 3). The further loss of Esox
sibiricus in Europe and success of morphologically modern
forms (e.g. Esox moldavicus, E. lucius, etc.) is assumed to be
linked to the changing global climate which underwent
a cooling phase after the Miocene Climate Optimum
(Mosbrugger et al. 2005; Ivanov et al. 2011). Increased sea-
sonality with a reduction in the number of warm weeks
meant that temperatures would have favoured species with
a wider temperature tolerance (Kovalchuk and Murray 2016).
The research of the ﬁrst author (O.K.) was supported by a grant for a visit to
Poland within the framework of cooperation between the Polish Academy
of Sciences and the National Academy of Sciences of Ukraine. W. Gornig is
acknowledged for making the photos of some studied specimens. We are
sincerely thankful to J. Divay and an anonymous reviewer for their con-
structive comments and advice. We also express our thanks to the editor
Gareth Dyke for his eﬀective management of our submission.
No potential conﬂict of interest was reported by the authors.
Oleksandr Kovalchuk http://orcid.org/0000-0002-9545-208X
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Figure 3. Miocene localities yielding the skeletal remains of Esox sibiricus:1–Sarybulak; 2–Holu; 3–Dsagzo Chairchan; 4–Javor; 5–Khirgis-Nur; 6–Tchono-
Chariak; 7–Popovo 3; 8–Lobkove (=Lobkovo); 9–Cherevychne (=Cherevichne); 10 –Verkhnya Krynitsya 2; 11 –Egorovka 2; 12 –Orikhivka (=Orekhovka); 13 –
Vynogradivka 1 (=Vinogradovka 1); 14 –Novo-Petrivka (=Novopetrovka, Kuchurgan); 15 –Bełchatów. Localities referenced by Sytchevskaya (1989) are indicated as
red circles, those mentioned in Kovalchuk et al. (2017)–as yellow circles, and Bełchatów –as a green circle.
6O. KOVALCHUK ET AL.
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Appendix Indeterminate ﬁsh remains (Teleostei indet.) from Bełchatów in the palaeoichthyological
collection of ZPALWr
ZPALWr. BM/69 –dentary fragment
ZPALWr. BM/70 –opercular fragment
ZPALWr. BM/56-58, BM/63 –broken abdominal centra
ZPALWr. BM/29-31, BM/48-55, BM/61-62, BM/64-66, BM/80-83, BM/90-99, BM/103, BM/105, BM/115-130, ВМ/140-149 –small bone fragments and debris
ZPALWr. BM/113 –bone breccia sample
8O. KOVALCHUK ET AL.