Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian
A new microvertebrate assemblage from Skelia Podil’ska 2 (Ternopil Oblast, the Dniester area, Ukraine) sheds light on faunal evolution during the Martonosha interglacial (Early Chibanian, Pleistocene). The fauna was discovered beneath a small rock shelter developed in the Shyrokyne alluvial sandston...
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| author | Popova, L. Veklych, Yu. Yanenko, V. Rekovets, L. Mironchuk, T. Stepanchuk, V. |
| author_facet | Popova, L. Veklych, Yu. Yanenko, V. Rekovets, L. Mironchuk, T. Stepanchuk, V. |
| author_institution_txt_mv | [
{
"author": "L. Popova",
"institution": "I. I. Schmalhausen Institute of Zoology NAS of Ukraine"
},
{
"author": "Yu. Veklych",
"institution": "Ukrainian Geological Research Institute of the State Enterprise «Ukrainian Geological Company»"
},
{
"author": "V. Yanenko",
"institution": "National Museum of Natural History, National Academy of Sciences of Ukraine"
},
{
"author": "L. Rekovets",
"institution": "Wrocław University of Environmental and Life Sciences, Poland"
},
{
"author": "T. Mironchuk",
"institution": "Taras Shevchenko National University of Kyiv, Ukraine"
},
{
"author": "V. Stepanchuk",
"institution": "Institute of Archaeology, NAS of Ukraine, Ukraine"
}
] |
| author_sort | Popova, L. |
| baseUrl_str | https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/oai |
| collection | OJS |
| datestamp_date | 2026-06-29T16:20:36Z |
| description | A new microvertebrate assemblage from Skelia Podil’ska 2 (Ternopil Oblast, the Dniester area, Ukraine) sheds light on faunal evolution during the Martonosha interglacial (Early Chibanian, Pleistocene). The fauna was discovered beneath a small rock shelter developed in the Shyrokyne alluvial sandstones. The section comprises alluvial and subaerial deposits of the Zbruch valley, and records several palaeoincisions of different ages. The oldest, Martonosha palaeoincision, formed a cliff face hosting a small rock shelter in which the fossils were recovered. Taphonomic analysis of the findings and their palaeogeomorphological context indicates that the bones accumulated due to owl activity. The fossil assemblage includes amphibians and rodents and is taxonomically and ecologically attributable to the Kolkotovian association of the Tiraspol fauna (MIS 17). The fauna is characterised by the absence of steppe and semi-desert species, and by a pronounced dominance of the subgenus Terricola voles, particularly Microtus (Terricola) arvalidens. Amphibian remains constitute an unusually high proportion of the assemblage. Ecologically, the assemblage reflects warm and highly humid climatic conditions typical of the Martonosha interglacial. At the same time, the scarcity of strictly forest-associated vertebrates suggests that closed forests were locally poorly developed. This discrepancy is attributed to shallow limestone and sandstone bedrock, which limited forest development despite favourable climatic conditions. |
| doi_str_mv | 10.15407/zoo2026.03.292 |
| first_indexed | 2026-06-30T01:00:28Z |
| format | Article |
| fulltext |
DOI 10.15407/zoo2026.03.292
UDC
FOSSIL VERTEBRATES OF SKELIA PODIL’SKA 2
(DNISTER AREA, UKRAINE): A DISTINCTIVE
INTERGLACIAL FAUNA OF THE EARLY CHIBANIAN
L. Popova 1, *, Yu. Veklych 2, V. Yanenko 3, L. Rekovets 4,
T. Mironchuk 5 & V. Stepanchuk 6, 7
1 I. I. Schmalhausen Institute of Zoology, NAS of Ukraine,
vul. B. Khmelnytskogo, 15,
Kyiv, 01054 Ukraine
E-mail: liliapopovalilia@gmail.com
2 Ukrainian Geological Research Institute of the State Enterprise
“Ukrainian Geological Company”,
78 A, Avtozavodska St., Kyiv, 04114 Ukraine
E-mail: veklych_um@ukr.net
3 National Museum of Natural History, National Academy of Sciences of Ukraine,
vul. B. Khmelnytskogo, 15, Kyiv, 01030 Ukraine
E-mail: ornithologist.ua@gmail.com
4 Wrocław University of Environmental and Life Sciences,
Curie-Skłodowskiej St. 42, 50-375 Wrocław, Poland
E-mail: leonid.rekovets@upwr.edu.pl
5 Taras Shevchenko National University of Kyiv,
Vasylkivska St., 90, Kyiv, 03022 Ukraine
E-mail: t.mironchuk@knu.ua
6 Institute of Archaeology, NAS of Ukraine, 12, V. Ivasiuka ave., Kyiv, 04210 Ukraine
7 I. Krypyakevych Institute of Ukrainian Studies, NAS of Ukraine,
Kozelnytska St., 4, Lviv, 79026 Ukraine
E-mail: VadimStepanchuk@gmail.com
L. Popova (https://0000-0001-5008-8715)
Yu. Veklych (https://orcid.org/0000-0002-7448-9342)
V. Yanenko (https://orcid.org/0000-0002-1241-1929)
L. Rekovets (https://orcid.org/0000-0001-9934-7095)
T. Mironchuk (https://0000-0002-5923-1028)
V. Stepanchuk (https://orcid.org/0000-0002-5476-2284)
urn:lsid:zoobank.org:pub:08D24CF0-48D0-4BE0-8796-BAAABEC1E2D8
Paleontology Zoodiversity, 60(3): 292–300, 2026
© Publisher Publishing House “Akademperiodyka” of the NAS of Ukraine, 2026. The article is
published under an open access license CC BY-NC-ND (https://creativecommons.org/licenses/
by-nc-nd/4.0/)
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
Fossil Vertebrates of Skelia Podil’ska 2 (Dnister Area, Ukraine): A Distinctive Interglacial
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
293
Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna
of the Early Chibanian. Popova, L., Veklych, Yu., Yanenko V., Rekovets, L., Mironchuk, T.,
Stepanchuk, V. — A new microvertebrate assemblage from Skelia Podil’ska 2 (Ternopil Oblast,
the Dniester area, Ukraine) sheds light on faunal evolution during the Martonosha interglacial
(Early Chibanian, Pleistocene). The fauna was discovered beneath a small rock shelter develo-
ped in the Shyrokyne alluvial sandstones. The section comprises alluvial and subaerial deposits
of the Zbruch valley, and records several palaeoincisions of different ages. The oldest, Martono-
sha palaeoincision, formed a cliff face hosting a small rock shelter in which the fossils were reco-
vered. Taphonomic analysis of the findings and their palaeogeomorphological context indicates
that the bones accumulated due to owl activity. The fossil assemblage includes amphibians and
rodents and is taxonomically and ecologically attributable to the Kolkotovian association of the
Tiraspol fauna (MIS 17). The fauna is characterised by the absence of steppe and semi-desert
species, and by a pronounced dominance of the subgenus Terricola voles, particularly Micro-
tus (Terricola) arvalidens. Amphibian remains constitute an unusually high proportion of the
assemblage. Ecologically, the assemblage reflects warm and highly humid climatic conditions
typical of the Martonosha interglacial. At the same time, the scarcity of strictly forest-associated
vertebrates suggests that closed forests were locally poorly developed. This discrepancy is attri-
buted to shallow limestone and sandstone bedrock, which limited forest development despite
favourable climatic conditions.
Ke y words : Dniester area, amphibians, small mammals, Terricola, Early Chibanian.
Introduction
The study of Pleistocene microvertebrates provides important insights into how va-
rious environmental factors contribute to shaping faunal communities. The Kolkoto-
vian association of the Tiraspol faunal complex is of a special interest in this respect.
The Kolkotovian fauna (Rekovets, 1994; Rekovets & Nadachowski, 1995; Krokhmal’
et al., 2021) existed in the interglacial Martonosha environment, characterised by the
expansion of forests, the richness of arboreal vegetation, and peak humidity (Gerasi-
menko, 2010). The Kolkotovian stage of the Ukrainian Pleistocene corresponds to the
Cromer II interglacial of Western Europe, MIS 17, i. e., 0.78–0.65 MY (Gerasimenko,
2010; Krokhmal’ et al., 2021). The correlation between the international scale and
Ukrainian regional stratigraphic units is described in Gradstein et al. (2020).
A characteristic trait of the Kolkotovian fauna is dominance of rodents pre-
ferring meadow and shrub habitats. Taxonomically, the dominant group consisted of
voles of subgenera Terricola Fatio 1876 and Microtus Shrank 1798 (Rekovets, 1994;
Rekovets & Nadachowski, 1995). Although such faunal composition is believed to
have been most characteristic of the late Martonosha time, similar assemblages mi-
ght have occurred among the Tiraspolian faunas until 0.45 MY (Rekovets & Nada-
chowski, 1995; Krokhmal’ et al., 2021; Krokhmal’ et al., 2023). Some Tiraspolian
small mammal associations included two species of Terricola: Microtus (T.) arva-
lidens (Kretzoi, 1958) and Microtus (T.) majori Thomas, 1906, a unique situation that
has never been repeated within the area of Ukraine since then (Rekovets & Nada-
chowski, 1995). Later, cooling and aridification led to a decrease in the occurrence
and diversity of Terricola. In the Late Pleistocene, Terricola vanished from the
Ukrainian area and had not reappeared until the Holocene (Popova, 2015).
The Kolkotovian fauna (the type locality is Kolkotova Balka III) was correlated
by Rekovets (1994) with a time interval of West Runton and Voigtstedt in the Wes-
L. Popova, Yu. Veklych, V. Yanenko, L. Rekovets, T. Mironchuk & V. Stepanchuk
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
294
tern European faunas, and on this basis, with MIS 17 (Krokhmal’ et al., 2021). Both
West- and East-European faunas of this time are characterised by their rich and
diverse composition indicating interglacial forest conditions; however, in contrast
to the Ukrainian Kolkotovian, there was no such marked increase in the share of
Terricola remains (Rekovets, 1994; Maul & Parfitt, 2010). Importantly, all currently
known Ukrainian Martonosha vertebrate taphocoenoses (Krokhmal’ et al., 2021)
were restricted to the Black Sea area, south of the reconstructed forest zone, and
largely retained steppe features. The discovery of a new taphocoenosis, Skelia Po-
dil’ska 2 (Early Chibanian, the Dnister area, Ternopil’ Oblast, Ukraine), for the first
time allows shedding light on the palaeoecological features of typical Martonosha
communities.
Geologica l and geomorphologica l sett ings
The Quaternary and Pre-Quaternary deposits in the vicinity of Skelia Podil’ska have
been repeatedly studied, as they provide a remarkably complete record of Neo-
gene-Quaternary sedimentation.
This area lies to the south of the most ancient terraces of the Dnister River
and belongs to the valley of its left tributary, the Zbruch River. Palaeosols and
loesses of the Shyrokyne-Martonosha-Sula stages (Late Calabrian-Early Chiba-
nian) are widely distributed in the adjacent area, while younger loess and soil
deposits, starting with the Lubny soil, may be exclusively seen in the palaeoinci-
sions. The exception is the Zavadivka soil, which is widely distributed there
(Fig. 1). This indicates a post-Sulian regional decrease in loess accumulation
rates, which, in turn, reflects large-scale lateral changes of the Quaternary sub-
aerial sedimentation (Veklych, 2018).
Krokhmal’ et al. (2009) revealed small mammal fauna in the alluvial depos-
its of the Zbruch River. Later, this fauna (Skelia Podil’ska 1) was attributed to the
earliest (Luzanivkian; Shyrokyne-Pryazovsk) phase of the Tiraspol complex and
MIS 20-21 (Krokhmal’ et al., 2021). Respectively, our section should be designat-
ed as Skelia Podil’ska 2.
The new locality, Skelia-Podil’ska 2, was revealed in 2017, in the course of archa-
eological prospecting conducted by the Lower Palaeolithic expedition of the Institu-
te of Archaeology of NASU, in the frame of “The earliest Palaeolithic sites of Ukraine
in the context of the initial peopling of Europe”, the project of the National Research
Fund of Ukraine (Stepanchuk et al., 2017; Popova et al., 2024; Stepanchuk & Vek-
lych, 2025).
The section of Skelia Podil’ska 2 is a wall of the quarry developed on the right
slope of the Zbruch valley. Pleistocene loams (15 m) ranging in age from the Holo-
cene to the Late Chibanian, overlie the Upper Calabrian gravel and medium-to-
coarse-grained sand and loose sandstone. The latter correspond to the sandstones
containing the small mammal fauna of Skelia Podil’ska 1 (Krokhmal’ et al., 2009).
These deposits are underlain by Badenian lithothamnion sandstones, clays, and
sands (Fig. 1).
A series of palaeoincisions is observed in the Pleistocene loam sequence (Fig. 1).
The most ancient Martonosha palaeoincision cuts the Shyrokyne hydromorphic soil
and alluvial sands and sandstones of the same age. The Martonosha slope soil (red-
Fossil Vertebrates of Skelia Podil’ska 2 (Dnister Area, Ukraine): A Distinctive Interglacial
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
295
dish-brown loams and brown dense loams up to 1.4 m in thickness) forms the lower
layer of the filling of this palaeoincision and dips steeply to the thalweg.
The age of the Skelia Podil’ska 2 microvertebrate fauna corresponds to the time
of this Martonosha incision. Fossil microvertebrates were found in the filling of a
minute rock shelter (with the maximal length of the cavity c. 0.3 m) in the loose
Shyrokyne alluvial sandstone beneath a denser sandstone layer of the same age. Sim-
ilar niches are frequently used by owls for nesting.
The rich-in-fossils filling of the niche (c. 2 kg) was collected into a plastic con-
tainer and screen-washed with a 1 mm mesh metal sieve.
Results and Discussion
The Skelia Podil’ska 2 microvertebrate fauna displays the characteristic features of
the Kolkotovian faunas: ecologically, by the absence of steppe species and domi-
nance of wetland forms, taxonomically, by dominance of Terricola, and biostrati-
graphically, by the evolutionary level of voles (Table 1).
Signs of digestion on vertebrate remains (Fig. 2, 6) indicate their accumulation
by avian or mammalian predators; while the size of the cavity with bones and its
Fig. 1. The outcrop of Skelia Podil’ska 2: 1 — Martonosha palaeoincision; 2 — Lubny palaeoinci-
sion and several Lubny palaeosols with loess interlayers; 3 — technogenic deposits of the quarry;
sh — sandstones and sands of the Shyrokyne alluvium; mr — reddish-brown loams of Martono-
sha soil; sl — the Sula loess; tl — Tyligul loess; zv — Zavadivka soil; Pg — Paleogene
L. Popova, Yu. Veklych, V. Yanenko, L. Rekovets, T. Mironchuk & V. Stepanchuk
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
296
position in the Martonosha relief (in the cliff, at least 4 m above the thalweg of the
valley) suggest owls as the most likely taphonomical agent. A complete absence of in
situ molars and virtually 100% of broken postcrania — unusual for bone accumula-
tion produced by predators — may be attributed to the increasing trampling effect
due to the heterogenous texture and granulometric composition of the floor of the
niche (lose sands with protruding denser sandstone, and more compact sandstone
ceiling debris — Fig. 1). Such composition should have increased mechanic grinding
of remains by trampling (Andrews, 1990). Relatively low total share of significantly
digested molars in the taphocoenoses may be due to the dominance of Terricola,
with its thick enamel on trailing ages of molar prisms. In contrast, the small sample
of Skelia Podil’ska 2 M. (M.) nivaloides consists of 50% moderate to heavily digested
molars (Table 1). Predatory digestion is known to affect prey molars differentially,
with voles showing higher degrees of digestion than murids (Demirel et al., 2011).
Here we observe a similar disparity for Microtus and Terricola.
Table 1 . Vertebrate remains from the Skelia Podil’ska 2 locality
Taxon
Number of identified remains
Minimum
number of
individualstotal
digestion remains
graded from
moderate to strong
juveniles
Anura indet 45 44 8
Bombina sp. 1 1 1
Hyla sp. 2 1 2
Rana cf. temporaria 4 4 3
Rana sp. 19 19 9
Pelobates cf. fuscus 5 5 2
Bufo sp. 16 12 4
Rana cf. arvalis 10 8 4
Pelophylax cf. ridibundus 9 8 5
Clethrionomys sp. 2 0 2 2
Microtus (Terricola) arvalidens 12 1 0 7
Microtus (Microtus) nivaloides Major, 1902 4 2 1 3
Microtus s. l. sp. 29 5 4 4
Fig. 2. Vole molars (m1) from Skelia Podil’ska 2: 1–4 — M. (Terricola) arvalidens; 5–7 —
M. (Microtus) nivaloides; 6 — moderately digested m1; 7 — juvenile m1. Scale bar 1 mm
Fossil Vertebrates of Skelia Podil’ska 2 (Dnister Area, Ukraine): A Distinctive Interglacial
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
297
The total absence of open-habitat species in the Skelia Podil’ska 2 is unique
among Pleistocene vertebrate faunas in Ukraine. Steppe, tundra or semi-desert spe-
cies are present in varying numbers in all other known localities; in most cases, they
are dominant (Rekovets, 1994).
On the other hand, Skelia Podil’ska 2 cannot be regarded as a typical forest
assemblage. A decisive argument for the zonal tree vegetation would be the presen-
ce of climbing tree-dwellers such as tree squirrels or glirids, which are absent in
Skelia Podil’ska 2. The bank vole Clethrionomys Tilesius, 1850, another rodent indi-
cator of the forest vegetation, is encountered in the fauna. The linkage of Clethrio-
nomys with the forest environment is not as definite as in the case of tree-dwelling
rodents since the bank vole may occur in pellet assemblages from the steppe zone
(Kondratenko et al., 2006). However, paleoecological studies demonstrate that its
occurrence has increased with climate warming and forest expansion (Rekovets,
1994). Still, the minor presence of Clethrionomys is a mere presence rather than the
dominance, which should be expected based on the other palaeoenvironmental
proxies (Gerasimenko, 2010). Amphibians of Skelia Podil’ska 2 do not belong to
strictly forest-associated species either; they are species of near-water habitats of the
forest-steppe or forest zone.
The absence or scarcity of forest species might have been partly explained by pred-
ator selectivity; however, in the case of Skelia Podil’ska 2, such an explanation is unlike-
ly. In particular, significant selectivity biases may be introduced by such predators as
the fox and the buzzard. The former is incapable of capturing tree-dwelling rodents.
The latter specialises in the prey from open habitats. Consequently, arboreal species
and Clethrionomys, a species that prefers dense vegetation, are typically underrepre-
sented in buzzard pellet assemblages (Andrews, 1990). Given the palaeogeomorpho-
logical settings at Skelia Podil’ska 2 point toward an owl as the primary taphonomic
agent, with owls being capable of capturing tree-dwelling species and the bank vole
(Andrews, 1990, Append. Fig. 3; comparison of owl pellets and data of trapping; and,
for the Ukrainian area, Kondratenko et al., 2006), the scarcity of forest fauna in Skelia
Podil’ska 2 requires another, palaeoecological explanation.
Ecological preferences of dominant small mammals of Skelia Podil’ska 2,
М. (M.) nivaloides and M. (Terricola) arvalidens are rather vague, which is quite
common for extinct species. Maul & Parfitt (2010) consider M. nivaloides to be asso-
ciated with open grasslands, while according to Pazonyi et al. (2018), this species was
confined to forested areas. Possibly, М. nivaloides was merely less environmentally
specialised compared to its likely descendants, such as M. arvalis and M. agrestis. In
any case, M. nivaloides belongs to relatively mesophilic rodents rather than to a
cold-tolerant or xerophilous ones.
Recent Terricola prefer meadow, shrub and forest habitats. Yet they rarely occur
in closed forest stands as wetlands are more preferable for them (Shenbrot & Abra-
mov, 1995). Ecological preferences of the extinct M. arvalidens are more diffuse.
Although an increased proportion of Terricola usually corresponds to warm and wet
stages of the Ukrainian Pleistocene, the species might have occurred in the steppe
faunas (Semybalka 2, the second Protopopovian association of the Tiraspolian com-
plex, MIS 19) and sometimes, even dominated in them (Tykhonivka 1 and Lysa
Gora, Tykhonovian and Nagirnian association of the Tiraspolian complex, MIS 16
L. Popova, Yu. Veklych, V. Yanenko, L. Rekovets, T. Mironchuk & V. Stepanchuk
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
298
and 12) (Rekovets & Nadachowski, 1995, Rekovets et al., 2014; Krokhmal’ et al.,
2021). Moreover, the mentioned localities belong to the dry Azov steppe, with two of
them, Tykhonivka 1 and Lysa Gora, assigned to the cold and dry stages of the Pleis-
tocene. Probably, M. arvalidens occupied intrazonal wet-valley habitats within those
zonal dry-steppe assemblages. Since all mentioned taphocoenoses of the Azov area
are alluvial, the high proportion of near-water and wetland species (including M.
arvalidens) among them is due to the local topography. In contrast, in the case of the
owl pellet assemblage of Skelia Podil’ska 2, the dominance of M. arvalidens is a signal
of climate humidification rather than a local phenomenon. Most Pleistocene Euro-
pean owl pellet assemblages are vole-dominated, but which species prevails is largely
determined by climate (Andrews, 1990). The increased total proportion of amphibi-
ans in Skelia Podilska 2, which significantly exceeds the levels typically observed in
different owl pellet assemblages (Andrews, 1990; Roulin & Dubey, 2013), shares the
same climatic explanation.
Therefore, the microvertebrate fauna from Skelia Podil’ska 2 reflects a warm, and
above all, highly humid climate. At the same time, this fauna does not provide a basis
for reconstructing a zonal forest landscape, which should necessarily be expected
under such conditions and is reconstructed on the basis of other palaeogeographical
proxies (Matviishyna et al., 2010). The absence or scarcity of typical forest-associated
vertebrates in the taphocoenosis may account for the local vegetation state. The
near-surface position of limestone and sandstone deposits in the Zbruch valley
during the Martonosha time (Fig. 1) is likely to have hindered the development of
the forest cover even under favourable climatic conditions.
The increase in M. (Terricola) arvalidens seems to be a regional peculiarity of the
Ukrainian Early Chibanian interglacials, particularly the Martonosha. This increase
is in line with some other features of these faunas, which are still not fully under-
stood. Thus, tree-dwelling rodents, which were common in the Pleistocene of Mol-
dova and Poland, are extremely scarce in the Pleistocene fossil record of Ukraine
(Rekovets, 1994) until the Holocene (Popova, 2015). Instead, Miomymus remains
were recorded (Rekovets, 1994). Among European dormice, this is the only genus
that avoids arboreal habitats. The structure of the Kolkotovian microvertebrate asso-
ciation definitely needs further study.
Conclusions
The newly discovered microvertebrate fauna from Skelia Podil’ska 2 (the Dnister
area, Ternopil' Oblast) corresponds to the Martonosha interglacial of the Early
Chibanian and is attributed to the Kolkotovian association of the Tiraspol faunal
complex (MIS 17).
Typical characters of the Kolkotovian (the decline of steppe species, dominance
of Terricola) are prominently manifested in Skelia Podil’ska 2.
Taphonomically, Skelia Podil’ska 2 is an owl pellet assemblage. The disparity in
the share of moderate to heavily digested molars has been registered between Micro-
tus and Terricola. This may be the consequence of differences in the enamel micro-
structure in voles of these subgenera.
Fossil Vertebrates of Skelia Podil’ska 2 (Dnister Area, Ukraine): A Distinctive Interglacial
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
299
The dominance of M. (Terricola) arvalidens and a high total proportion of am-
phibian remains in Skelia Podil’ska 2 indicate a significant humidification of the cli-
mate; in contrast, the absence or scarcity of strictly forest-associated vertebrates may
be attributed to specific geological settings of the area (a near-surface position of
limestone and sandstone deposits, which prevented a complete forestation despite
the climatic conditions of the Martonosha time).
Acknowledgements. The research of LP was carried out as part of the research
project ‘Formation of highly integrated complex structures in evolution and devel-
opment of animals’, supported by the I. I. Schmalhausen Institute of Zoology, NAS of
Ukraine, and was partly financed by the Krzysztof Skubiszewski Foundation. Field-
work was supported by the State Fund for Fundamental Research of Ukraine under
Project No. F77/91-2017 (0117U003691) (VSt).
REFERENCES
Andrews, P. 1990. Owls, caves and fossils: predation, preservation and accumulation of small
mammal bones in caves, with an analysis of the Pleistocene cave faunas from Westbury-sub-
Mendip, Somerset, UK. University of Chicago Press, Chicago, 1–230. https://books.google.
com/books/about/Owls_Caves_and_Fossils.html?id=GZaMNt55oT8C
Demirel, A., Andrews, P., Yalçınkaya, I. & Ersoy, A. 2011. The taphonomy and palaeoenvi-
ronmental implications of the small mammals from Karain Cave, Turkey. Journal of Archa-
eological Science, 38 (11), 3048–3059. https://doi.org/10.1016/j.jas.2011.07.003
Gerasimenko, N. P. 2010. Correlation of short-termed stages of the Pleistocene according to pa-
laeolandscape data. In: Matviishyna, Z. N., Gerasimenko N. P., Perederii V.I., Bragin, A. M.,
Ivchenko, A. S., Karmazynenko, S. P., Nahirnyi, V. M. & Parkhomenko, O. D. Spatio-tem-
poral correlation of palaeogeographic conditions of the Quaternary in Ukraine. Naukova
dumka, Kyiv, 104–128 [In Ukrainian].
Gradstein, F. M., Ogg, J. G., Schmitz, M. D. & Ogg, G. M. 2020. The Geologic Time Scale 2020 (2). Else-
vier. 1–1349. https://www.sciencedirect.com/book/9780128243602/geologic-time-scale-2020
Kondratenko, O., Kuznetsov, V. & Timoshenkov, V. 2006. Peculiarities of diet of the long-ea-
red owl (Asio otus) in the Donets-and-Don and Azov steppes. Proceedings of Theriological
School, 7, 77–79.
Krokhmal’, O., Komar, M. & Prylypko, S. 2009. Results of palaeontological examination of allu-
vial deposits of the Eopleistocene terrace of the Zbruch river near Skala Podil’ska village
(Ternopol’ Region). In: Bogutski, A. Most ancient loesses of Podilla and Pokuttia. Collection
of materials of XVI Ukrainian-Polish scientific conference. L’viv University Editorial Center,
L’viv, 166–168 [In Ukrainian].
Krokhmal’, O., Rekovets, L. & Kovalchuk, O. 2021. An updated biochronology of Ukrainian
small mammal faunas of the past 1.8 million years based on voles (Rodentia, Arvicolidae): a
review. Boreas, 50, 619–630. https://doi.org/10.1111/bor.12503
Krokhmal’, O., Rekovets, L. & Kovalchuk, O. 2023. Biochronological scheme of the Qua-
ternary of the south of Eastern Europe and its substantiation based on arvicoline teeth
morphometrics. Quaternary International, 674–675, 5–17. https://doi.org/10.1016/j.
quaint.2023.05.012
Matviishyna, Z. N., Gerasimenko N. P., Perederii V. I., Bragin, A. M., Ivchenko, A. S., Karma-
zynenko, S. P., Nahirnyi, V. M. & Parkhomenko, O. D.. 2010. Spatio-temporal correlation
of palaeogeographic conditions of the Quaternary in Ukraine. Naukova dumka, Kyiv, 1–191
[In Ukrainian].
L. Popova, Yu. Veklych, V. Yanenko, L. Rekovets, T. Mironchuk & V. Stepanchuk
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 3
300
Maul, L. C. & Parfitt, S. A. 2010. Micromammals from the 1995 Mammoth Excavation at West
Runton, Norfolk, UK: Morphometric data, biostratigraphy and taxonomic reappraisal. Qua-
ternary international, 228, (1–2), 91–115. https://doi.org/10.1016/j.quaint.2009.01.005
Pazonyi, P., Virág, A., Podani, J. & Pálfy, J. 2018. Microtus (Microtus) nivaloides from the
Somssich Hill 2 site (southern Hungary): An Early Pleistocene forerunner of modern ‘true’
Microtus voles revealed by morphometric analyses. Quaternary International, 481, 1–14.
https://doi.org/10.1016/j.quaint.2017.07.019
Popova, L. V. 2015. Small mammal fauna as an evidence of environmental dynamics in the Ho-
locene of Ukrainian area. Quaternary international, 357, 82–92. https://doi.org/10.1016/j.
quaint.2014.10.021
Popova, L., Veklych, Y., & Stepanchuk, V. 2024. Fossil small mammals of Skelia Podil’ska 2
(Dnister area, Ukraine): A distinctive interglacial fauna of the Early Chibanian. Abstract
Book of the International conference Climate Change in Europe: Impact, Risks and Mitiga-
tion (CCE2024), Vatra Dornei, Suceava county, Romania, 10–13 October 2024. Geoconcept
Journal, 5 (1), 44–47.
Rekovets, L. I. 1994. Small mammals of the Antropogene of the south of Eastern Europe. Nauko-
va dumka, Kyiv, 1–370 [In Russian].
Rekovets, L. & Nadachowski, A. 1995. Pleistocene voles (Arvicolidae) of the Ukraine. Paleonto-
logia i evolució, 28, 145–246.
Rekovets, L., Čermak, S., Kovalchuk, O., Prisyazhniuk, V., & Nowakowski, D. 2014. Vertebrates
from the middle pleistocene locality Lysa Gora 1 in Ukraine. Quaternary International, 326,
481–491. https://doi.org/10.1016/j.quaint.2013.10.016
Roulin, A. & Dubey, S. 2013. Amphibians in the diet of European Barn Owls. Bird Study, 60 (2),
264–269. https://doi.org/10.1080/00063657.2013.767307
Shenbrot, G. I. & Abramov, A.V., 1995. Taxonomy and ecology of shrub voles (Terricola). In:
Wilson, D. E. & Reeder, D. M., eds. Mammal Species of the World: A Taxonomic and Geo-
graphic Reference. 2nd ed. Washington, DC: Smithsonian Institution Press. https://www.
departments.bucknell.edu/biology/resources/msw3/
Stepanchuk, V. M., Bakhmutov, V. G., Matviishyna, Zh. M., Dmytruk, Yu. M., Karmazynen-
ko, S. P., Veklych, Yu. M., Tsvelykh, O. M., Kovalchuk, O. M., Vetrov, D. O., Nezdoliy, O. I.
& Vetrov, V. S. 2017. Report on SRP 0117U003691 “The oldest Paleolithic sites of Ukraine
in the context of the initial colonization of Europe”. Institute of Archeology of NASU, Kyiv,
2017. Deposited in UkrINTEI, Kyiv, 1–37 [In Ukrainian].
Stepanchuk, V., Veklych, Y. 2025. Lower Palaeolithic surveys in Western Ukraine (2017): Archae-
ological, Geomorphological, and Geological aspects. Materials and research on archaeology of
Carpathian and Volhynia, 29, 109–147. https://doi.org/10.33402/mdapv.2025-29-109-147.
Veklych, Yu. M. 2018. Geoeolian morpholithogenesis and methodological aspects of its research.
UkrDGRI, Kyiv, 1–253 [In Ukrainian].
Received 14 January 2026
Accepted 30 June 2026
|
| id | oai:ojs.akademperiodyka.org.ua:article-919 |
| institution | Zoodiversity |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2026-06-30T01:00:28Z |
| publishDate | 2026 |
| publisher | Publishing House "Akademperiodyka" of the National Academy of Sciences of Ukraine |
| record_format | ojs |
| resource_txt_mv | ojsakademperiodykaorgua/0e/99486d4ce644695171615b6fd482440e.pdf |
| spelling | oai:ojs.akademperiodyka.org.ua:article-9192026-06-29T16:20:36Z Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian Popova, L. Veklych, Yu. Yanenko, V. Rekovets, L. Mironchuk, T. Stepanchuk, V. Dniester area amphibians small mammals Terricola Early Chibanian A new microvertebrate assemblage from Skelia Podil’ska 2 (Ternopil Oblast, the Dniester area, Ukraine) sheds light on faunal evolution during the Martonosha interglacial (Early Chibanian, Pleistocene). The fauna was discovered beneath a small rock shelter developed in the Shyrokyne alluvial sandstones. The section comprises alluvial and subaerial deposits of the Zbruch valley, and records several palaeoincisions of different ages. The oldest, Martonosha palaeoincision, formed a cliff face hosting a small rock shelter in which the fossils were recovered. Taphonomic analysis of the findings and their palaeogeomorphological context indicates that the bones accumulated due to owl activity. The fossil assemblage includes amphibians and rodents and is taxonomically and ecologically attributable to the Kolkotovian association of the Tiraspol fauna (MIS 17). The fauna is characterised by the absence of steppe and semi-desert species, and by a pronounced dominance of the subgenus Terricola voles, particularly Microtus (Terricola) arvalidens. Amphibian remains constitute an unusually high proportion of the assemblage. Ecologically, the assemblage reflects warm and highly humid climatic conditions typical of the Martonosha interglacial. At the same time, the scarcity of strictly forest-associated vertebrates suggests that closed forests were locally poorly developed. This discrepancy is attributed to shallow limestone and sandstone bedrock, which limited forest development despite favourable climatic conditions. Publishing House "Akademperiodyka" of the National Academy of Sciences of Ukraine 2026-04-27 Article Article application/pdf https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/919 10.15407/zoo2026.03.292 Zoodiversity; Vol. 60 No. 3 (2026): Zoodiversity Zoodiversity (Vestnik Zoologii); Том 60 № 3 (2026): Zoodiversity 2707-7268 2707-725X 10.15407/zoo2026.03 en https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/919/399 Copyright (c) 2026 L. POPOVA, Y. Veklych, V. Yanenko, L. Rekovets, T. Mironchuk, V. STEPANCHUK |
| spellingShingle | Popova, L. Veklych, Yu. Yanenko, V. Rekovets, L. Mironchuk, T. Stepanchuk, V. Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian |
| title | Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian |
| title_full | Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian |
| title_fullStr | Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian |
| title_full_unstemmed | Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian |
| title_short | Fossil vertebrates of Skelia Podil’ska 2 (Dnister area, Ukraine): a distinctive interglacial fauna of the Early Chibanian |
| title_sort | fossil vertebrates of skelia podil’ska 2 (dnister area, ukraine): a distinctive interglacial fauna of the early chibanian |
| topic_facet | Dniester area amphibians small mammals Terricola Early Chibanian |
| url | https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/919 |
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