Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea)
A paleogeographic reconstruction of Quaternary continental water bodies in Ukraine based on ostracods requires the data necessary for correlating the stages of development of water bodies in Eastern and Western Europe to be generalised, and synchronised with the general climatic trend of the Qua...
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| author | Dykan , N. I. |
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| author_institution_txt_mv | [
{
"author": "N. I. Dykan ",
"institution": "Institute of Geological Sciences of NAS of Ukraine, Ukraine ",
"orcid": "0000-0001-6631-7041"
}
] |
| author_orcid_str_mv | 0000-0001-6631-7041 |
| author_sort | Dykan , N. I. |
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| container_title | Zoodiversity (Vestnik Zoologii) |
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| datestamp_date | 2026-08-26T09:51:17Z |
| description |
A paleogeographic reconstruction of Quaternary continental water bodies in Ukraine based on ostracods requires the data necessary for correlating the stages of development of water bodies in Eastern and Western Europe to be generalised, and synchronised with the general climatic trend of the Quaternary. A variety of methods and analyses were employed, including actualistic, quantitative modification, taphonomic, statistical, population, ecological, zoogeographical and biostratigraphic approaches. The detailed reconstruction of river and lake paleo-biotopes (type of biotope, degree of trophicity, water depth of biotope, m; water temperature , °C) in the paleo-valleys of the Zbruch, Southern Bug, Psol and Dnipro rivers during the Early to Middle Pleistocene (Calabrian – Holsteinian) has been conducted. A comparative analysis of the development of continental water bodies in Ukraine, Lithuania, Belarus and Central Russia during the Middle Pleistocene (Holsteinian) was conducted based on the results of studying freshwater ostracods of Eastern Europe. Correlating fluctuations (increases/decreases) in the water levels of paleo-water bodies located at considerable distances from each other reveals a general trend in water level changes caused by Interglacial Holsteinian climatic oscillations. It also reveals asynchrony (the partial displacement of positive and negative peaks relative to each other) and different water level fluctuation amplitudes caused by geographical zoning of the territory. According to the ostracods, the Holsteinian interglacial period had three warm stages (temperate stages I, II and III) and two cold stages (cold stages I and II). In southern water bodies (Ukraine), three stages of warming (temperate stages I, II and III) are recorded by an increase in water temperature. The first and third stages of warming (temperate stages I and III) are also recorded by an increase in water level. The second period of warming (temperate stage II) occurred during the first prolonged cold stage (cold stage I), which is indicated by rising water levels in northern bodies of water and only by an increase in water temperature in southern water bodies. Based on ostracods the timing of the formation and dynamics of a paleo-landslide in the paleo-valley of the Zbruch River during the Early Pleistocene (Calabrian) have been reconstructed.
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| doi_str_mv | 10.15407/zoo2026.04.342 |
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| first_indexed | 2026-08-27T01:00:40Z |
| format | Article |
| fulltext |
DOI 10.15407/zoo2026.04.342
UDC 565.33:551.8(477)"624"
PALEOGEOGRAPHIC RECONSTRUCTION
OF THE CONTINENTAL WATER BODIES OF UKRAINE
IN THE EARLY-MIDDLE PLEISTOCENE BASED
ON OSTRACODS (ARTHROPODA, CRUSTACEA)
N. I. Dykan
Institute of Geological Sciences of NAS of Ukraine,
vul. O. Gonchara, 55-b, Kyiv, 01054 Ukraine
E-mail: natalidykan@gmail.com
N. Dykan (https://orcid.org/0000-0001-6631-7041)
urn:lsid:zoobank.org:pub:4FA34009-6BC8-4C73-8C83-941DF323EC66
Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Mid-
dle Pleistocene based on ostracods (Arthropoda, Crustacea). Dykan, N. I. — A paleogeo-
graphic reconstruction of Quaternary continental water bodies in Ukraine based on ostracods
requires the data necessary for correlating the stages of development of water bodies in Eastern
and Western Europe to be generalised, and synchronised with the general climatic trend of the
Quaternary. A variety of methods and analyses were employed, including actualistic, quantita-
tive modification, taphonomic, statistical, population, ecological, zoogeographical and biostrati-
graphic approaches. The detailed reconstruction of river and lake paleo-biotopes (type of biot-
ope, degree of trophicity, water depth of biotope, m; water temperature, °C) in the paleo-valleys
of the Zbruch, Southern Bug, Psol and Dnipro rivers during the Early to Middle Pleistocene
(Calabrian – Holsteinian) has been conducted. A comparative analysis of the development of
continental water bodies in Ukraine, Lithuania, Belarus and Central Russia during the Middle
Pleistocene (Holsteinian) was conducted based on the results of studying freshwater ostracods of
Eastern Europe. Correlating fluctuations (increases/decreases) in the water levels of paleo-water
bodies located at considerable distances from each other reveals a general trend in water level
changes caused by Interglacial Holsteinian climatic oscillations. It also reveals asynchrony (the
partial displacement of positive and negative peaks relative to each other) and different water
level fluctuation amplitudes caused by geographical zoning of the territory. According to the
ostracods, the Holsteinian interglacial period had three warm stages (temperate stages I, II and
III) and two cold stages (cold stages I and II). In southern water bodies (Ukraine), three stages
of warming (temperate stages I, II and III) are recorded by an increase in water temperature. The
first and third stages of warming (temperate stages I and III) are also recorded by an increase in
Paleontology Zoodiversity, 60(4): 342–382, 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. 4
Paleogeographic Reconstruction of the Continental Water Bodies of Ukraine
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 4
343
water level. The second period of warming (temperate stage II) occurred during the first pro-
longed cold stage (cold stage I), which is indicated by rising water levels in northern bodies of
water and only by an increase in water temperature in southern water bodies. Based on ostracods
the timing of the formation and dynamics of a paleo-landslide in the paleo-valley of the Zbruch
River during the Early Pleistocene (Calabrian) have been reconstructed
Key words: freshwater ostracods, ecology, Eastern Europe, paleoclimate, Holsteinian
Introduction
Benthic ostracods are an important source of data for paleogeographic reconstruc-
tions of ancient (continental, marine) paleobasins. A wide range of ostracod toler-
ance combined with a narrow ecological adaptation (stenobiont) determine their
widespread geographical distribution and existence in various water bodies. Ostra-
cods occupy different positions in food chains and have a certain ecological special-
ization, and the species composition of assemblages of ostracods reflects the peculi-
arities of their habitat in biotopes (water bodies). Fossil ostracods are numerous,
sufficiently well preserved for species identification, and predominantly autochtho-
nous in Quaternary deposits. In addition, the structure of populations (adults, lar-
vae, females, males) is preserved in fossil ostracods, which contains information
about all, even minor, changes in the aquatic environment. The analysis of the con-
ditions of existence of recent ostracods and their extrapolation to ancient environ-
ment are based on the principle of actualism, which implies the similarity of modern
and past geological events. At the same time, it is essential to take into account the
natural patterns of functioning of recent biota, the cause-and-effect series of ecolog-
ical laws and rules of functioning of biosystems that determine the existence and
spread of recent organisms (Reimers, 1991, 1994). 97% of Quaternary ostracods
continue to exist in modern ecosystems of continental water bodies and the Black
Sea (Dykan, 2005, 2006, 2012, 2016).
Quaternary freshwater ostracods in Ukraine have been studied fragmentarily
(Matoshko et al., 2002, Dykan, 2006). The first data included a description of the
species composition of ostracods from alluvial deposits of the rivers of Western
Ukraine (rivers Danube, Prut, Dniester; Schneider & Konstantinova, 1966; Nega-
daev-Nikonov, 1974). Since the 1990s, studies of river terraces in Western Ukraine
(rivers Zbruch, Prut, Southern Bug), Northern and Central Ukraine (rivers Dnipro,
Norin, Desna, Sula, Ros, Vorskla), and Southern Ukraine (rivers Danube, Berda,
Molochna) new data on the biostratigraphy of Lower Pleistocene-Holocene alluvial
deposits, paleogeographic reconstructions of river and lake paleo-water bodies, and
the ecology of recent freshwater ostracods have been obtained (Dykan, 1994 a, b,
1995, 1996, 2001 a, b, 2003, 2008, 2014, 2015) (Fig. 1). Paleogeographic reconstruc-
tion of Quaternary continental water bodies in Ukraine based on ostracods requires
generalization of numerous data necessary for correlating the stages of development
of water bodies in Eastern and Western Europe and synchronization with the gener-
al climatic trend of the Quaternary.
The aim of the study was to investigate the stages of development of freshwater
ostracods in the continental (river and lake) water bodies of Ukraine during the
Pleistocene, the correlation with the continental paleo-water bodies of Eastern Eu-
N. I. Dykan
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 4
344
rope, and, at the final stage of analytical generalization, the reconstruction of the
climatic cyclicality of the Interglacial Holsteinian in the Holarctic zone (48–60° N).
Material and Methods
The actual material was the author’s collection of fossil and recent ostracods from
natural outcrops of continental water bodies of Ukraine (rivers, lakes), collected dur-
ing 1981–2022 (collection no. 2061 is housed at NMNH of NAS of Ukraine; collec-
tions no. 2567, 2589, 3000 is housed at the Department of Quaternary Geology, IGN
of NAS of Ukraine, Kyiv).
A wide range of methods and analyses (actualistical, modified quantitative, tapho-
nomical, statistical, population, ecological, zoogeographical, biostratigraphical) were
used in paleogeographic reconstructions of Quaternary continental water bodies based
on ostracods (Dykan, 2005, 2016, 2023, 2026).. Paleontological data for other fauna
groups, results of paleopedological analysis, and paleomagnetic dating were also taken
into account. This allowed us to obtain a wide range of data on fossil freshwater ostra-
cods, make biostratigraphic conclusions about the geological age of alluvial deposits and
paleogeographic conclusions regarding the conditions of ostracods existence in the Ear-
ly-Middle Pleistocene. The biostratigraphic division of Quaternary alluvial deposits in
Ukraine was carried out in accordance with SKU (2012); chronostratigraphic correlation
with general subdivisions (Series, Subseries, Stages) and regional units (Ukrainian Loess
Plain Stages, North West European Stages) and their substages are given according to
GTS2020 (Gradstein et.al., 2020) and Global chronostratigraphical correlation table for
the 2.7 million years (Cohen & Gibbard, 2019, 2022).
Results
Lower-Middle Pleistocene alluvial deposits of river terraces in Western and Central
Ukraine (rivers Zbruch, Southern Bug, Psel, Dnipro) contain ostracods diverse in
genus (15 genera) and species (32 species) composition (Tables 1–2 and fig. 6, 15 see
in https://doi.org/10.5281/zenodo.21918941; Paleontological tables 3–6, here and
further tables see in the Appendix).
Alluvial deposits of the Lower Pleistocene (Gelasian-Calabrian)
Geological section “Scala Podilska” (profiles A, B, C), the Zbruch River (Middle
Dniester River basin), VII terrace of the Dniester River basin, the southwestern part of
Podolian Plateau (270 m a. s. l.), the village of Burdyakivtsi (construction quarry),
Chortkiv District, Ternopil Region, Western Ukraine, coordinates 48°50′ N, 26°10′ E
(Boguckyja et al., 2009) (Fig. 1, here and further pictures see in the Appendix).
In the central part of the quarry, to the right of the landslide (profile A), the fol-
lowing are exposed (from bottom to top): Paleozoic bedrock (Silurian: limestones,
visible thickness 10.1 m, layer 1); Neogene (Middle Miocene, Langhian/Badenian;
marine deposits: lithotamnium limestones, sandstones, clays, sands; total thickness
20 m, layer 2), Quaternary (Lower Pleistocene, Tiglian/Beregovian-Eburonian/Be-
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razanian, alluvial deposits: gravelly horizon, sands, loams, thickness 6.8 m, layers
3–6; Lower Pleistocene-Holocene, Waalian/Kryzhanivian-Eemian/Prylukian, sub-
aerial deposits: loess, paleosols, thickness 11 m, layers 7, 9; redeposited deposits of
Serravalian/Sarmatian, deluvial deposits: loams, thickness 5.8 m, layer 8) (Dykan,
2008; Boguckyja et al., 2009) (Fig. 2).
Alluvial deposits lie on the lithotamnium limestones of Langhian/Badenian,
overlain by the subaerial deposits of Lower Pleistocene (Waalian/Kryzhanivian). Al-
luvium represented by channel facies (profile A) and floodplain facies (profiles B, C)
of one alluvial cycle of the Zbruch River paleovalley (Fig. 3). The Lower Pleistocene
age of the alluvium was determined according to paleomagnetic (TL age), paleonto-
logical and palynological data (mollusks, plant pollen) (Dykan, 2008; Boguckyja et al.,
2009). Among ostracods, the species Limnocythere tuberculata Negadaev, 1957 (de-
scribed from Lower Pleistocene, Gelazian; Balkan Peninsula; Krstić, 1985) and Prion-
ocypris zenkery (Chyzer, 1858) (described from Lower Pleistocene, Calabrian, Moldo-
va; Negadaev-Nikonov, 1974) have biostratigraphic significance. Other species (Cyp-
ria candonaeformis (Schweyer, 1949), Ilyocypris gibba (Ramdohr, 1808), I. bradyi Sars,
1890) are transient in Quaternary deposits. The basal horizon (redeposited gravel of
silicon, sandstone, limestone; without paleontological remains; layer 3) lies at the bot-
tom of alluvial deposits. The time of formation of the basal alluvium refers to “...the
period of development of the primary river network of the Carpathian foreland..., to
the first stage of local erosion and dissection... of Carpathian gravel deposits and base-
ment rocks” and coincides with Gelasian (Tiglian/Beregovian; Boguckyja et al., 2009:
201–202). The accumulation time of channel and floodplain alluvium facies (layers
4–6) correlates with the Lower Calabrian (Eburonian/Berezanian).
Alluvial deposits contain numerous remains of freshwater ostracods of autoch-
thonous burial (Ilyocypris, Prionocypris, Limnocythere, Candona, Cypria) (Fig. 4;
Table 1). The accumulation of channel alluvium (Profile A) occurred under condi-
tions of weakening deep erosion processes, a gradual increase in channel depth and
water temperature (Fig. 5). A few ostracods existed in shallow (0.3–0.5 m) riverbed
biotopes on sandy substrate, with a water temperature of +(18–20) °C (layer 4).
These are reophile and cryophile Il. bradyi (single adults and larvae; typical habitats
of recent representatives of the species are flowing water bodies, cold springs, a wa-
ter depth up to 0.1–0.5 m, temperature pessimum more than +(18–19) °С; Bron-
shtein, 1947; Kovalenko. 1976; Meisch, 2000; Fuhrmann, 2012); cryophile P. zenk-
ery (single adults; recent representatives exist at water depths of up to 0.3–0.5 m,
water temperature not exceeding +20 °C; Kovalenko, 1976; Meisch, 2000; Fuhr-
mann, 2012); eurydaphic L. tuberculata (single adults). The species composition of
ostracods changed with the subsequent increase in water depth to 3 m and an in-
crease in water temperature to +(23–24) °C (layer 5). Thermophile, phytophile and
deep-water stenobat C. candonaeformis appeared and became dominant (typical
biotope is slow-flowing rivers, lakes, oxbows; depth 3.0–6.0 m, water temperature
+(23–24) ºС, dense aquatic vegetation; Kovalenko 1987). Il. bradyi had the high
population density (adults, larvae), and the presence of the phytophile P. zenkery
(single adults; typical biotopes are slow streams and cold springs with dense vegeta-
tion; Kovalenko, 1976; Meisch, 2000) indicates the presence of cold springs and the
development of dense bottom vegetation.
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Two temporary floodplain water basins were reconstructed in the Zbruch pale-
ovalley (Calabrian, Eburonian/Berezanian) (layer 6). The ostracod association was
represented by typical inhabitants of lakes and oxbows. The first floodplain water
body (lake; Profile B) was deep (up to 4 m) and warm (+23–25 ºС). Thermo-phy-
tophile C. candonaeformis was dominated. The few thermophile Il. gibba appeared
(this species exists in modern water bodies at water depth of up to 1.0–1.5 m and
water temperatures of up to +25 ºС; Kovalenko, 1976, 1988). The second floodplain
water body was shallow (up to 1.5 m; profile C), with cold springs and dense aquatic
vegetation. Two species, small numbers of cold-loving phytophiles Il. bradyi and
P. zenkery, existed in the marshes around cold springs (Bronshtein, 1947; Kovalen-
ko, 1976).
Reconstruct ion of the dynamics and t iming of pa leolandsl ide for-
mat ion in the Zbruch pa leoval ley (Early Pleistocene, Calabrian).
Deposits of uncertain origin (layer 8, thickness 5.8 m; Figs 2, 3) lie in the lower part
of the subaerial deposits, between the loess horizon (Menapian/Ilichivian) and the
paleosoil horizon (Cromerian complex, Interglacial I-II/Martonoshian). They are
represented by greenish-gray loam, homogeneous, with layers and lenses of detritus,
numerous paleontological remains (ostracods, unidentified fish otoliths, bryozoans,
hedgehog needles, foraminifera, Miocene (re-deposited) marine and brackishwater
mollusks); “... these enigmatic greenish-grey loams” (Bogutsky et al., 2009: 179). The
loam have fluvial-paludic genesis and “...the functioning of a shallow lake with weak
currents, fed with spring water and small streams... lake periodically dried out, and
paludal deposition occurred...” (Bogutsky et al., 2009: 179). The time of loam accu-
mulation is comparable to Bivelian/Shyrokinian (Calabrian, Lower Pleistocene). At
the same time, this hypothesis does not explain the presence of marine autochtho-
nous/synchronous ostracods, as well as other marine organisms in the oryctoceno-
sis, according to which the age of the layer 8 loam is determined as Lower Serraval-
lian/Sarmatian (Middle Miocene). Autochthonous ostracods account for 26% of the
total number of marine Sarmatian species (Aurila sarmaticа, Haplocytheridea daci-
ca, Hemicytheria omphalodes, Loxoconcha kochi). Synchronous ostracods account
for 36% of the total number of Sarmatian species (Loxoconcha ornatа, Cytheridea
hungarica, Aurila cicatricosа, Pterygocythereis jonesii, Cnestocythere truncata, Loxo-
concha rhombovalis, Cushmanidea lithodomoides, Callistocythere molesta) (Dykan,
2008) (Fig. 4).
According to the analysis of the geological structure of the Podolian Plateau, the
presence of a thick layer of Sarmatian deposits (5.8 m, layer 8) between the subaerial
horizons of Quaternary deposits (layers 7 and 9) is, in the author’s opinion, the result
of an ancient landslide process in the paleovalley of the Zbruch River.
The alluvium is cut into the thickness of the plateau (270 m a. s. l.), composed in
the upper part of Sarmatian deposits with a thickness of about 30–40 m. The bottom
of alluviam lies at the dates 230–235 m a. s. l. and 30–35 m above the water level of
the Zbruch River. That is, the close transfer of fossil remains was possible either by
direct erosion of Sarmatian deposits or by deluvial processes on the slopes of the
paleovalley. Pedological and sedimentological analysis of subaerial deposits of layer
7 (partially preserved Menapian/Ilichivian loess horizon and clearly eroded Waalian/
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Kryzhanivian illuvial humus soil) also indicates a significant local erosion during the
accumulation of loess-soil deposits (Boguckyja et al., 2009). The active development
of modern landslide processes in the studied area (groundwater discharge from nu-
merous sources occurs through Silurian and Baden limestones) is additional confir-
mation of the possible landslide nature of the Sarmatian deposits of layer 8 (Fig. 2).
Conclusion. The accumulation of alluvial deposits in the paleovalley of the
Zbruch River was replaced by the accumulation of subaerial deposits. A suffosion
slip of part of the slope composed of Sarmatian deposits occurred during the accu-
mulation of the loam-soil complex of layer 7 (Waalian/Menapian) along the dis-
placement surface, which was formed by plastic Sarmatian clays. The loam-soil ho-
rizons of layer 7 (Waalian/Kryzhanivian, Menapian/Ilichivian) were partially eroded
during the displacement of the landslide block, therefore the time of the paleoland-
slide is determined by the following stage, Bavelian/Shyrokinian. The accumulation
of subaerial deposits (buried soil, Cromerian complex, Interglacial I-II/Martonoshi-
an) on the surface of the landslide body continued after the landslide (Figs 2, 4).
Alluvial deposits of the Middle Pleistocene (Chibanian)
Geological section “Medschybisch” (Medschybisch 1 “Reindeer” profile,
Medschybisch “A” profile; Lower Paleolithic archaeological site, Oldovan culture of
ancient man, Stepanchuk et al., 2014), the Southern Bug River, IV floodplain terrace,
Podolian Plateau, town of Medschybisch, Letychiv District, Khmelnytskyi Region,
Western Ukraine, coordinates 49°35′40′′ N, 27°42′23′′ E (Figs 1, 6).
In the geological section “Medschybisch” the following are exposed (from bottom to
top): Archean bedrock (granites), Neogene marine deposits (Serravallian/Lower Sarma-
tian; gravel-pebble horizon, sands; visible thickness 1.2 m), Quaternary deposits (Middle
Pleistocene, alluvial deposits: sands, loams, loamy sands, thickness 2.05–4.4 м; Middle
Pleistocene–Holocene, subaerial deposits: loess, paleosols, thickness 3.1–5.0 m) (Matvi-
yishyna & Karmazynenko, 2014; Dykan, 2014). Alluvial deposits lie on the weathering
crust of Archean granites (Medschybisch 1 “Reindeer” profile) and marine coastal
sandy-gravel-pebble deposits of the Serravalian/Lower Sarmatian (Medschybisch “A”
profile), are overlain by paleosols of the Eemian/Kaydakian horizon (Fig. 7).
The age of the alluvium (Middle Pleistocene, Chebanian) was determined ac-
cording to paleontological (mollusks, fish, small and large theriofauna), paleoped-
ological, palynological, and archaeological (Lower Paleolithic hominid artifacts)
methods (Medschybisch 1 “Reindeer” profile); archaeological method (Lower
Paleolithic hominid artifacts) (Medschybisch “A” profile) (Stepanchuk, 2014). Os-
tracods are represented by transitional species from the Neogene (Middle Mio-
cene) to modern water bodies and have no biostratigraphic significance (Table 2).
Channel facies (Medschybisch 1 “Reindeer” profile) and floodplain facies
(Medschybisch 1 “Olenyachiy” and Medschybisch “A” profiles) were studied for
ostracods. Ostracods are numerous and diverse in genus (Cypria, Cyclocypris,
Cypridopsis, Cyprinotus, Candona, Typhlocypris, Ilyocypris) and species (Il. brad-
yi, C. candonaeformis, Cyclocypris ovum (Jurine, 1820), Cypridopsis vidua
(Müller, 1776), Cyprinotus salinus (Brady, 1862), Candona candida (O. Müller,
1776), Typhlocypris rostrata (Brady and Norman, 1889). All ostracod remains
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have an autochthonous type of burial (Fig. 8). Two types of floodplain water
bodies have been reconstructed: a periodically drained, shallow water body
(Medschybisch “A” profile) and a semi-flowing lake of the oxbow type (Medschy-
bisch 1 “Reindeer” profile) (Fig. 9).
Medschybisch “A”prof i le. A continuous series of Cromerian/Martonoshi-
an–Saalian/Dnieperian floodplain facies of the Middle Pleistocene has been studied
in the Medschybisch “A” profile (Figs 7, 9, 10). The floodplain alluvium of the warm
stage of the Interglacial I-II, Cromerian complex/Martonoshian (layer 20, thickness
0.55 m) is represented interlayers of sandy loam, loam, red-brown, black, strongly
oxidized sands; at the base by sands dark gray, clayey with pebbles and limestone
fragments; paleontological remains include ostracods and archaeological artifacts.
Ostracods were represented only by single adult individuals of the rheophilic and
cryophile Il. bradyi (recent representatives of the species are few in number at depths
of more than 0.5 m and temperatures above +(18–19) ºС. The reconstructed biotope
had a water depth slightly more than, and a water temperature above +(18–19) ºC, as
evidenced by the low population density and absence of larvae in Il. bradyi.
The floodplain alluvium of the cold stage of the Glacial b, Cromerian complex/
Sulian (layer 19, thickness 0.15 m) is represented by interlayers of gray and red-
dish-brown sands, with pebbles, boulders, and limestone fragments; paleontological
remains are ostracods. The species Il. bradyi was the dominant species and has the
high population density (only adults; optimal habitat water depth of recent repre-
sentatives is the coastal zone up to 0.5 m, water temperature is around +11–12 ºС,
and aquatic vegetation). A small number (individual specimens) of eurythermic
(close to thermophilic) C. salinus (Brady, 1862) (the recent representatives live at
water depths of up to 0.30 m, the temperature range is +11–26 °C; Kovalenko 1976)
and еurythermic, close to cold-loving Il. gibba (water depth from 0.3 m to 1.0–1.5 m,
temperature range +4–26 °C; Dесkkеr, 1979; Kovalenko, 1976) have colonized the
biotope. The high population density of Il. bradyi, the appearance of the cold-loving
Il. gibba and the shallow-water phytophilе C. salinus indicate that the floodplain wa-
ter body has become shallower (0.1–0.5 m) and colder (+11–12 ºС), and bottom
vegetation developed.
The floodplain alluvium of the warm stage of the Interglacial III–IV, Cromerian
complex/Lubenian (layer 18, thickness 0.60 m) is represented interlayers of gray-
brown and red-brown sands, with lenses of gray clayey sand, and sandy-pebble inter-
layers of eroded Sarmatian deposits; paleontological remains include ostracods and
archaeological artifacts (bone fragments and remains of ancient campfires). Ostra-
cods are represented by Il. gibba (average population density) and the few C. salinus.
The disappearance of the shallow-water сryophile Il. bradyi and the increase in the
population density of Il. gibba (it occurs as single individuals at +20–25 °C) indicate a
slight increase in depth (up to 1.0 m) and an increase in water temperature to +20 °C.
The alluvium of the cold stage of the Elsterian/Tiligulian (layer 17, thickness
0.15 m) is represented by a coastal channel facies (gray and light gray sands). Ostra-
cods are absent, indicating the significant cooling and temporary disappearance of
the floodplain water body.
The floodplain alluvium of the warm stage of the Holsteinian/Zavadivian (layer
16 a, thickness 0.50 m) is represented interlayers of dark gray sandy loams and gray,
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red-brown sands; paleontological remains include ostracods and archaeological ar-
tifacts (remains of ancient campfires). Ostracods are represented only by single lar-
vae of Il. bradyi, indicating a decrease in the water depth of the biotope to 0.5 m and
a decrease in water temperature to +(18–19) ºС.
The alluvium of the cold stage of the Warthe Drenthe, Saalian/Dnieperian (layer
14 a, thickness 0.2 m) is represented of light gray, fine-grained sands; paleontological
remains are animal bones) does not contain ostracod remains, which indicates the
significant cooling and disappearance of shallow water bodies.
C onclusion. The floodplain water body was cold (water temperature ranged
from +11 to 25 °C), and cold springs were developed in the coastal zone. The water
body was shallow (depth ranged from 0.1 to 1.0 m) and ceased to exist during cold
periods (Elsterian/Tiligulian, Warthe Drenthe Saalian/Dnieperian) (Figs 9, 10).
During warm periods (Lubenian/Cromeria, Zavadivian/Holsteinian), hominid set-
tlements were located on the water body shore (Oldowan culture of the Holsteinian/
Zavadivian era; Stepanchuk et al., 2014).
Medschybisch 1 “Reindeer” prof i le. Floodplain facies (Holsteinian/Zavadivi-
an) and channel facies (Warthe Drenthe, Saalian/Dnieperian) of alluvium were studied
in the Medschybisch 1 “Reindeer” profile (Fig. 11). Floodplain facies (layers 15–16,
total thickness 1.5 m) lie on the eroded surface of alluvial deposits is represented by
light brown, iron-rich loams, with pebbles and gravel (thickness 0.1 m). Floodplain
facies contain numerous paleontological remains. There are ostracods, mollusks
(Viviparus, Theodoxus, Lithoglyphus, Unio), vertebrate bones (deer, bear, southern ele-
phant, beaver), archaeological artifacts (stone tools and bone fragments with signs of
human processing) (Matviishyna & Karmazinenko, 2014; Gozhik et al., 2014; Stepan-
chuk et al., 2014). The genus and species composition of ostracods is close to the ostra-
cod associations of the Medschybisch “A” profile. Ostracods are numerous and diverse
in genus (Cypria, Cyclocypris, Cypridopsis, Candona, Typhlocypris) and species (C. can-
donaeformis, C. ovum, C. vidua, C. candida, T. rostrata). Оstracods have autochthonous
and synchronous types of burial (Dykan, 2014) (Fig. 9).
Floodplain deposits of layer 16 (thickness 1.1 m) are represented by brown,
iron-rich loams and sandy loams, with gravel and pebbles, with layers of numerous
horizontally oriented valves and shells of Unio spp. (Fig. 12). Ostracods are repre-
sented by only one species, the limno-thermo-phytophilе C. candonaeformis (recent
representatives of species are typical inhabitants of lakes and oxbows, living at water
depths of 3.0–6.0 m, at a water temperature of +23–24 °C). The reconstructed biot-
ope had a water depth of 3–6 m, water temperature of +23–24 °C, and developed
dense aquatic vegetation, as evidenced by the optimal development of Cypria can-
donaeformis (high density and composition of the population, where 82% were
adults and 8% were larvae of various ontogenetic stages) (Fig. 9).
Floodplain deposits of layer 15 are represented by brown iron-rich loams (thick-
ness 0.4 m, paleontological remains include ostracods, mollusks, freshwater fish, an-
imal bones; archaeological artifacts; Matviishyna & Karmazinenko, 2014; Gozhik et
al., 2014; Stepanchuk et al., 2014; Kovalchuk & Rekovets, 2014; Dykan, 2014).
The species composition of ostracods in the bottom layer 15 (layer 15 a) indi-
cates a decrease in a water depth and water temperature in the floodplain water body.
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The biotope was colonized by a few species of similar ecological specialization – lim-
nobians and phytophiles, resistant to a wide range of temperatures C. ovum, C.vidua,
T. rostrata. The thermophile C. candonaeformis dominated in terms of abundance,
but its population structure changed (ratio of aduals/larvae, 52 to 48%, respectively),
indicating a deterioration in the species’ habitat conditions. The water depth range in
which recent representatives of species C. ovum (shallow drying ponds, coastal areas
up to 2.0 m), C. vidua (up to 3.0 m), T. rostrata (up to 1.0 m) exist indicates a de-
crease in the water depth of the biotope to 1–2 m. Eurythermic, close to thermo-
philes species C. ovum (temperature range of recent representatives +14–26 °C in
ponds with spring water inflow, springs, lakes) and T. rostrata (up to +26 °C), as well
as eurythermic, close to cold-loving C. vidua (water temperature +8–26 °C; found
throughout the year, with lower numbers in winter than in summer) are an indicator
of a slight decrease in water temperature to +20–23 °C.
At the end of the Holsteinian/Zavadivian stage (layer 15 b), the floodplain lake
became shallow (about 1 m) and cold (up to +16–20 °C), and a new composition of
ostracod communities formed. The stenothermic, cold-loving phytophilе C. can-
dida of medium population density (adults, larvae) appeared (the optimal condi-
tions for the existence of recent representatives of the species are shallow standing
water bodies and shallow water bodies with cold springs, water temperature +10–
16 °C, egg laying occurs at +4–10 °C, upper temperature range +20 °C). The
deep-water thermophile C. candonaeformis reduced population density by half.
T. rostrata increased its population density by 2 times (recent representatives are
numerous in shallow waters), C. ovum by 4.5 times. The relatively deep-water spe-
cies C. vidua disappeared (Fig. 9).
The alluvium of the cold Warthe Drenthe, Saalian/Dnieperian stage is repre-
sented by the coastal zone facies of the riverbed (layers 13–14, thickness 2.05 m).
There are light gray, fine-grained sands, sandy loams, clays; paleontological remains
are represented ostracods, mollusks, animal bones and archaeological artifacts (re-
mains of ancient hearths). The shallow biotope of the coastal zone of the riverbed
(layer 14, thickness 0.7 m) is represented by gray sands with light gray interlayers
and fine grain size. The dominance and threefold increase in the abundance of the
stenothermic cold-loving species C. candida, as well as a threefold decrease in the
abundance of the thermophile C. candonaeformis, are clear indicators of a further
decrease in water temperature to +(10–16) °C and water depth of the biotope from
1.0 m to 0.5 m.
At the end of the Saalian/Dnieperian stage (layer 13, thickness 0.5–1.35 m; inter-
layers light gray, iron-rich sand, with layers of clay and sand), the depth of the biot-
ope was less than 0.5 m, and the water temperature was less than +10 °C. Only one
species, T. rostrata, existed in the biotope in small numbers (isolated individuals).
Subsequently, this section of the reservoir ceased to exist, and the ostracods disap-
peared (Fig. 9).
Conclusion. At the beginning of its existence, the floodplain semi-flowing
lake was deep (up to 3–6 m) and warm (+23–24 °C), with undisturbed bottom sedi-
ments after their accumulation (layers 15, 16). The autochthonous type of burial of
fossil ostracods, as well as the taxonomic signs of fossil mollusk remains (subhori-
zontal and horizontal occurrence of shells and valves of Unio spp., good preservation
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and presence of unopened shells, and the absence of sorting of remains by size con-
firm the calm hydrodynamic regime in the lake (Fig. 12). Subsequently, there was a
gradual decrease in the depth of the water body (up to 0.3 m) and a decrease in water
temperature (to less than +10 °C). During the Glacial stage of Warthe Drenthe,
Saalian/Dnieperian, the water body disappeared and the accumulation of subaerial
deposits began (Kaydakian/Eemian) (Fig. 13).
Geological sect ion “Hun’ky” , the Psel River (left tributary of the Dnipro Riv-
er), IV floodplain terrace, Dnipro Lowland, north of the village of Hun’ky, Kremenchug
District, Poltava Region, Central Ukraine, coordinates 49°14’ N, 33°33 'E (Fig. 1).
In the river outcrop, the following are exposed (from the water level to top):
Quaternary deposits (Middle Pleistocene, Holsteinian/Zavadivian, alluvial deposits:
sands, loams, loamy sands, clays, lake marl, thickness 12–16.0 m; Middle Pleisto-
cene, Warthe Drenthe, Saalian/Dnieperian, glacial deposits, moraine, thickness
3.2 m); Middle Pleistocene–Holocene, subaerial deposits: loess, palesoils, thickness
10 m) (Fig. 14).
The age of the alluvial channel facies (Elsrerian/Tiligilian, layer 1) was deter-
mined according to mollusks (Gozhik, 2006). The age of floodplain facies (Holstein-
ian/Zavadivian, layers 2–10) was determined by geological methods based on the
occurrence of floodplain alluvium below the moraine of the Dnieperian horizon
(Saalian/Dnieperian), as well as paleontological methods according to ostracods,
mollusks, and small theriofauna (Dykan, 1994 a; Gozhik, 2006; Krokhmal & Reko-
vets, 2010). The index species Interglacial Holsteinian/Zavadivian is the freshwater
species L. tuberculata (Holsteinian/Saalian is the upper boundary of the stratigraph-
ic range of this species). The geographical barrier to the eastward expansion of
L. tuberculata was the Ural Mountains, and the ecological factor leading to its extinc-
tion in continental waters of Europe was the onset of the Glacial Saalian/Dniepero-
vian (Middle Pleistocene) (Dykan, 2001 a, b, 2003, 2008) (Fig. 15).
The channel facies of alluvium (layer 1, thickness 6.0–7.0 m) are represented by
white, yellow, fine-grained sands, obliquely-horizontally layered and does not con-
tain ostracod remains. Floodplain alluvium facies (layers 2–10, thickness 6.0–9 m)
contain ostracods of autochthonous burial type, diverse in genus (11 genera) and
species (23 species) composition (Fig. 16).
In the bottom of floodplain deposits (layer 2, thickness 0.3 m; interlayers of
blue-gray and ocher-colored, fine-grained sands), ostracods are represented by
12 species, mainly limnobionts and potamobionts. The dominant species were the
deep-water thermophile Сурriа candonaeformis (water depth of 3.0–6.0 m) and the
eurythermic, close to the thermophiles Сyclocypris laevis (О. F. Müller, 1785) (in-
habiting coastal and open areas up to 3–3.5 m; optimum temperature of +20–23 °C),
as well as the cold-loving, deep-water species Darwinula stevensoni (Brady et Rob-
ertson, 1870) (high population density at a water temperature of +11–15 °C and a
water depth of 6–10 m). Low population densities (single individuals) had euryther-
mic, cold-loving P. zenkeri (temperature range from +4 °C to no more than +22 °C),
Il.bradyi (normal development up to +12 °C; unfavorable conditions are tempera-
tures above +18–19 °C and water depths from 0.5 m to 4.0 m), Il. gibba (temperature
range from +4 to 19.5 °C, simple individuals are found at temperatures above
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+20 °C), Candona (E.) balatonica Daday, 1894 (temperature range from +10.5 °C to
17 °C, water depth of 0.2–50 m), Candona (E.) protzi Нartwіg, 1898 (temperature
range from +1°C to 15 °C, water depth up to 5.0 m). Other species were also few in
number. There are eurythermic Неrреtосурris rерtаns (Baird, 1835), (water depth of
0.2–5.0 m); eurythermic, close to thermophiles Candona (C.) elongata (Schweyer,
1949) (water depth of 1–4 m) and Limnocythere inopinata (Baird, 1843) (tempera-
ture range from +4 °C to 28 °C, water depth of 0.7–15 m), euryhedaphic L. tubercu-
lata. Analysis of the ecology ostracods (cold-loving species accounted for 50% of the
total composition of the association) and dynamics of ostracod population density
allows us to reconstruct the floodplain water body as relatively deep (1.5–3.5 m) and
cold (water temperature +15–20 °C), with cold springs at the bottom at the begin-
ning of its existence. The edaphic factor was unfavorable for the development of
cold-loving Candona, active mud-dwelling organisms, in the water body (popula-
tions of species were represented by single adults and numerous larvae) (Fig. 17).
Deposits of layer 3 (thickness 0.3 m) are represented by light gray loams, iron-
rich in the roof, with fragments of small shells. The species diversity of ostracods has
decreased by half. Deep-water species C. (C.) elongata and С. candonaeformis have
disappeared, as well as some cold-loving species (D. stevensoni, C. (E.) balatonica,
C. (E.) protzi, P. zenkeri). The number of heat-loving C. laevis has doubled (temper-
ature optimum of species +20–23 °C); the eurythermal, close to thermophiles L. in-
opinata has increased seven times. The decrease in the number of cold-loving and
deep-water species, as well as the increase in the number of heat-loving species, in-
dicate an increase in water temperature up to +(20–23) °С and a decrease in the
water depth up to 1 m. The phytophile Candona (С) neglecta Sars, 1887 and Candona
(С) angulata G. Müller, 1900 appeared, while the cold-loving phytophile Il. bradyi
increased its population density threefold (a typical inhabitant of cold springs and
marshes with high population densities), indicating the presence of cold springs and
the development of dense aquatic vegetation (Fig. 17).
The deposits of layer 4 (thickness 0.2 m) are represented by ochre-yellow sand
with small shell fragments and dark gray sand in the roof. Significant changes in the
species composition of ostracods occurred, with species diversity doubling (13 spe-
cies) (Fig. 22). Phytophile С. candonaeformis appeared and became dominant. Shal-
low-water spesies, Ilyocypris botniensis Kovalenko, 1976 (depth of existence no more
than 1–1.5 m) and phytophile Турhlocypris compressa (Kosh, 1837) (high population
density at depths of less than 1 m) appeared, as well as recurrent cold-loving species
C. (Е.) protzi (adults are found at water temperatures no more than +14–15 °С) and
C. (Е.) bаlаtonica (temperature range from +10.5 °С to 17 °С). The population den-
sity of C. laevis has declined to a few individuals (the species is small in number at a
water temperature of +14.8 °C and a water depth of more than 1 m). The relatively
deep-water Турhlocypris pratinensis (Hartwig, 1901) has disappeared (water depth of
existence from 1 m to 3–5 m). Unfavorable factors for deep-water Сandona, active
mud dwellers, were sandy substrate and shallow biotope (the number of larvae in
populations doubled). These changes in the species composition of ostracods, the
appearance of shallow-water and cold-loving species, and the decline in the abun-
dance of C. laevis indicate an increase in water level up to 1.5 m and a decrease in
water temperature up to +(10–15) °С, dense aquatic vegetation (Fig. 17).
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The deposits of layer 5 (thickness 0.8 m) are represented by dark gray, homoge-
neous, fine-grained, clayey sands with mollusk shells. There have been minor chang-
es in the composition of the association of ostracods (12 species). Cold-loving eury-
batiс species C. (С.) angulata (temperature range of +10–17 °С, water depth range
from 0.3 m to 30 m), stenobatic Candona (C.) iliensis Mandelstam, 1963 (depth 3 m),
relatively eurybatic Т. compressa (recent representatives found at a water depth of up
to 8 m, optimal water depth up to 1 m; temperature range of +6–26 °С, adults found
at temperature not lower than +18 °С), shallow-water Il. bоtniensis (water depth up
to 1–1.5 m) disappeared from the biotope. Relatively deep-water species have ap-
peared. There are the abundant D. stevensoni (optimal depth of 1.6–10 m), the few
species of C. (C.) elongata (water depth of 1–4 m), Н. rерtаns (water depth of 0.2–
5 m), as well as the cold-loving inhabitant of springs and streams with dense vegeta-
tion P. zenkeri (temperature range from +4 °C to no more than +20–22 °C). Cold-lov-
ing, more shallow-water species Il. bradyi (small numbers at water temperature
above +18–19 °C, water depth of 0.1–4 m) and Il. gibba (simple individuals occur at
temperature above +20 °C) reduced population densities. Cold-loving species C. (Е.)
protzi and C. (Е.) bаlаtonica continued to exist in the biotype, suggesting a wider
range of tolerance of these species to water temperature. The species C. laevis (water
depth up to 3–3.5 m; optimum temperature of +20–23 °C) increased population
density by 20 times and was dominant in the ostracod association. The accumulation
of clayey homogeneous sands of the floodplain facies occurred under conditions of
further water depth increase to 1.5–4 m and a slight increase in water temperature to
+19–23 °C (due to the continued existence of cold-loving species). Cold-loving spe-
cies Candоnа and phytophiles P. zenkeri, Н. rерtаns lived around cold springs among
dense aquatic vegetation (Fig. 17).
Deposits of layer 6 (thickness 0.4 m) are represented by gray loam, iron-rich at
the bottom, with mollusk shells. The species diversity of ostracods decreased to
9 species (Fig. 22). The number of Candona species decreased, while the single
cold-loving, relatively deep-water species C. (E.) protzi increased its population den-
sity sixfold (upper temperature threshold no more than +14–15 °C; water depth up
to 5 m, lower water depth threshold not defined) and became the dominant species
in the ostracod association. The cold-loving Il. brady had a population of medium
density (there is range between the temperature optimum, +12 °C, and pessimum,
+18 °C for recent representatives of the species). Eurythermic, close to the thermo-
philes C. laevis (simple individuals are found at a water temperature of +14.8 °C and
a water depth of more than 1 m), as well as the thermophile and deep-water C. can-
donaeformis (water temperature of +23–24 °С, water depth of 3–5 m) have reduced
their numbers to a few individuals. Cold-loving species Cyclocypris globosa (Sars,
1862) (temperature range of recent representatives of +4–18 °С) and Eucypris sp.
(most species of the genus Eucypris are cold-loving) have appeared. These changes in
the species composition and density of ostracod populations indicate a slight de-
crease in the water depth of the water body to 1–3 m and a sharp decrease in water
temperature to +12–15 °С (Fig. 17).
The alluvial facies (layer 7, thickness 1.4 m) are represented by lake marl (gitya)
dark gray to black, carbonaceous, with a large amount of organic matter, covered
with light gray marl with numerous remains of mollusks and small mammals;
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Gozhik, 2006; Krokhmal & Rekovets, 2010). Ostracods had low species diversity
(7 species in the bottom of layer 7a, two species in the upper part of layer 7 b). The
biocenosis was formed mainly from few, cold-loving and shallow-water species
Il. bradyi, Il. gibba, Typhlocypris insculpta (G.Müller, 1900) (adults are found at +18–
23 °С, water depth up to 0.4 m), Cyprinotus vassoevichi Schneider in Bodina, 1961
(recent representatives of the genus Cyprinotus inhabit the coastal zone of water
bodies up to 0.5 m), the relatively eurybiont Т. compressa, and the relatively deep-wa-
ter C. (C.) iliensis. With the gradual shallowing of the water body and a decrease in
water temperature, the number of C. (С.) iliensis larvae decreased, and the thermo-
philes and deep-water species of the genera Limnocythere, Cypria and Candоnа dis-
appeared. In the lake marl roof (layer 7 b), ostracods were represented by only two
species, namely the few Denticulocythere dorsotuberculata (Negadaev, 1957) and
Il. bradyi with medium population densities (more numerous than in the ostracod
association of layer 6). The reconstructed waater depth of the water body was up to
0.1 m, and the water temperature was +11–12 °С (Fig. 17).
The clay layer above (layer 8, thickness 0.1 m) does not contain any ostracod
remains.
The deposits of layer 9 (thickness 0.4 m) are represented by gray sandy loam
with a micro-layered subhorizontal texture at the bottom, with mollusk shells.
The species diversity of ostracods increased to five species. Thermophilic,
deep-water, few (single individuals) species C. candonaeformis, L. inopinata, as
well as relatively eurythermic, shallow-water Il. gibba (single individuals at
+20 °С and above, water depth up to 1.0–1.5 m) appeared. The reconstructed
biotope was warm (the water temperature rose to +18–26 °С), and the water
depth increased to 1.5 m (Fig. 17).
During the accumulation of layer 10 deposits (thickness 0.4 m; interlayering of
light gray silt and gray, fine-grained, finely subhorizontal layered sand), rapid shal-
lowing and a decrease in water temperature occurred, and ostracods disappeared
from the biotopes.
Rapid shallowing and a decrease in water temperature occurred during the ac-
cumulation of deposits of layer 10 (interlayering of light gray siltstone and gray sand,
fine-grained, with subhorizontal thin layering, 0.4 m thick), which led to the disap-
pearance of ostracods. Later, the floodplain water body ceased to exist.
Conclusion. The floodplain water body that existed in the paleovalley of the
Psel River during the Zavadivian/Holsteinian had an unstable hydrodynamic regime
with periodic fluctuations in water depth and temperature (Figs 17, 18):
1. At the beginning of its existence, the water body was relatively deep (1.5–
3.5 m) and cold (water temperature did not exceed +15–20 °С), with sparse vegeta-
tion or none at all, with outflows of cold spring waters (layer 2).
2. The subsequent shallowing of the water body (to 0.1–1 m) was accompanied
by an increase in water temperature to +(20–23) °C, the dense aquatic vegetation
developed, cold springs existed (layer 3).
3. The next stage was a gradual increase in water depth from 1–1.5 m (layer
4) to 4 m (layer 5), accompanied initially by a significant decrease in water tem-
perature to +10–15 ºС (layer 4), and then an increase in water temperature to
+19–23 °C (layer 5).
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4. At the next stage, the depth of the reservoir decreased from 1–3 m (layer 6) to
0.1 m (layer 7 b), and the water temperature decreased from +(12–15) °С (layer 6) to
+11–12 °С, layer 7 b).
5. At the final stage of its existence, the water body became deeper (0.5–1.5 m)
and warmer (water temperature rose to +18–26 °С, layer 9).
6. The subsequent rapid shallowing and decrease in water temperature led to the
disappearance of ostracods (layer 10), and the disappearance of the floodplain water
body. Subsequently, the floodplain ceased to exist.
Geological sect ion “Melnyky”, left bank of the Dnipro River, IV floodplain
terrace, Dnipro Lowland, Melnyky village, Tscherkassy District, Tscherkassy Region,
Central Ukraine, coordinates 49°18' N, 32°17' E (Fig. 1).
The river outcrops, the following are exposed (from the water level to top): Qua-
ternary deposits (Middle Pleistocene, Holsteinian/Zavadivian: alluvial deposits, lay-
ers 1–2, total thickness 1,8 m: channel facies, layer 1, sands, thickness 1.4 m; flood-
plain-type soil, layer 2, thickness 0.4 m); lake deposits, layers 3–8, siltstone, total
thickness 10.2 m; Middle Pleistocene, Warthe Drenthe, Saalian/Dnieperian, glacial
deposits, layer 9, moraine, thickness 11.4 m; Holocene, subaerial deposits, layer 10,
modern soil, thickness 0.3 m) (Fig. 19). Ostracods are fragmentarily distributed in
the vertical section of alluvial and lake deposits, absent in alluvial deposits (layers
1–2) and lake deposits (layers 3, 8) and numerous and diverse (8 genera, 10 species)
in lake deposits (layers 4–7), (Dykan, 1994 b) (Fig. 20).
The age of lake facies (Holsteinian/Zavadivian) was determined by geological
methods based on the occurrence lake deposits (layers 3–8) of below the moraine of
the Dnieperian horizon (Saalian/Dnieperian, layer 9). In the Melnyky and Hun’ky
sections, common species account for 50%. Recent representatives of fossil ostra-
cods in the Melnyky section account for 30% of the total number of species, in the
Hun'ky section – 13%, which allows us to consider the time of accumulation of lake
sediments in the Melnyky section to be younger than the alluvial deposits of Hun’ky
section and assign them to the second half of the Holsteinian/Zavadivian (Tables 1,
2). The considerable thickness of lake deposits (10.2 m), homogeneity of lithological
composition and textural features (siltstones are homogeneous, thinly layered) indi-
cate the stability of sedimentation conditions in the paleo-water body and its consid-
erable water depth. The “lake” type of ostracods, where 50% are limnobionts, typical
inhabitants of lakes and other permanent standing water bodies (Physocypria fadee-
vi Dubowsky, 1927, C. (E.) balatonica, Н. rерtаns, С. laevis, Т. compressa), allows us
to reconstruct the paleo-water body as a large drainless lake. Fossil ostracods and
other flora and fauna remains are absent in the lower part of the lake deposits (layer
3, thickness 5.8 m), indicating extremely unfavorable conditions for the develop-
ment of organic life in the lake at the beginning of its existence (low water tempera-
ture, great depth of the reservoir, absence of aquatic vegetation.
The first ostracods to colonize the water body (layer 4) were the cold-loving
species C. (E.) balatonica (inhabiting large standing water bodies, water temperature
of +10.5–17 °С, water depth of 0.2–5 m, optimal habitat being the coastal shallow
zone less than 1 m). The presence of only a few larvae in the populations indicates
that conditions are still unfavorable for the ontogenesis of C. (E.) balatonica (the
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water temperature was slightly above +10 °С, and the depth of the biotope was about
5 m) (Fig. 21).
In the overlying deposits (layer 5), the diversity of ostracods increased signifi-
cantly (5 genera, 6 species). The water body was colonized by the dominant, thermo-
phile and phytophile Т. compressa (adults are found at water temperature not lower
than +18ºC and are numerous in shallow areas of standing water bodies up to 1 m
deep); a few (single adult) species of С. lаеvis (single individuals at water depths
more than 1 m and water temperature of +14.8 °С) and the phytophile Н. rерtаns
(water depth of 0.2–5 m). The conditions for C. (E.) balatonica remained unfavorable
and numerous larvae prevailed in the populations. Changes in the species composi-
tion of ostracods and the presence of hytophiles indicate the appearance of aquatic
vegetation, an increase in water temperature (up to +18–20 °C) and a decrease in
depth to (at least) 1 m (Fig. 21).
The next stage of the paleolake’s development (layer 6) was favorable and opti-
mal for ostracods, which increased their genus and species diversity (8 genera,
10 species). Numerous phytophile P. fadeevi and few (single individuals) eury-eda-
phile L. inopinata (water depth of 0.7 –15 m) appeared. The species Т. соmрrеssа and
C. (E.) balatonica increased the population density (adults and larvae) threefold,
С. laevis (numerous at a water temperature of +20–23 °C) doubled, Paralimnocythere
originalis (Negadaev, 1965) increased fivefold, and Н. rерtаns increased 13-fold.
These changes among ostracods indicate an increase in water temperature to +(20–
23) °C and a subsequent decrease in water depth to 0.7–15 m, as well as the develop-
ment of dense bottom vegetation (Fig. 21).
In the overlying deposits (layer 7) the species L. inopinata and P. fadeevі disap-
peared. The population density of Т. соmрrеssа and C. (E.) balatonica decreased by
6 times, the species P. originalis by 2.5 times, and the species Н. rерtаns by 0.5 times.
С. laevis remained the dominant species in the ostracod association. These changes
in the association were caused by further shallowing of the lake up to 0.2 m and a
decrease in water temperature up to +(15–17) °C. There are no ostracods in the over-
lying sediments of layer 8 (Fig. 21).
Conclusion. The trend of lake development in the Dnipro paleovalley during the
second half of the Interglacial Holsteinian/Zavadivian was as follows (Figs 21, 22):
1. The paleo-water body was a large, drainless, deep lake. At the early stage of its
existence, the lake was very cold (less than +10 °С) and deep (significantly more than
5 m), without aquatic vegetation, undeveloped or extremely poor in organic life (os-
tracods did not inhabit the lake either). The few cold-loving Candona were the first
ostracods to colonize lake biotopes (the water depth of the reconstructed biotope
was about 5 m), with a water temperature slightly above +10 °C; layer 4);
2. The depth of the water body continued to decrease and amounted up to (at
least) 1 m, the water temperature rose up to +18–20 °C, and aquatic vegetation de-
veloped. The diversity of ostracods increased (layer 5);
3. The next stage was optimal for the development of ostracods. The water body
was warm (the water temperature rose to +20–23 °C), shallow (the water depth de-
creased to 0.7–1 m), with dense aquatic vegetation. Various and numerous ostra-
cods, limnobionts, and phytophiles gradually colonized the water body (layer 6);
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4. Further shallowing up to 0.2 m and a sharp decrease in water temperature up
to +15–17 °C became pessimistic factors for the existence of ostracods in shallow,
cold water bodies with sparse aquatic vegetation and led to a reduction in population
size and a decrease in ostracod species diversity (layer 7);
5. At the final stage of its existence, the paleo-lake became very cold and shallow,
and the bottom vegetation disappeared, leading to the extinction of all ostracods. At
the end of the Holsteinian/Zavadivian period, the water body disappeared.
Discussion
The Interglacial Holsteinian/Zavadivian was the most optimal period in the develop-
ment of freshwater ostracods in Ukrainian water bodies during the Quaternary. A
certain regularity in the change of ecological groups of fossil ostracods is recorded in
the vertical sections of alluvial deposits of the Dniester, Southern Bug, Dnipro, and
Psel rivers, which correlates with the periodicity of fluctuations of abiotic parameters
(water depth, water temperature) in river paleo-water bodies in the Holsteinian/Za-
vadivian. Reconstruction of water depth based on quantitative and qualitative anal-
ysis of fossil ostracods in adjacent territories of Eastern Europe (Lithuania, geologi-
cal section “Nyaravai”; Belarus, sections “Matveev Rov,” “Kolodyazhny Rov”; Russia,
“Likhvin” section) in Holsteinian/Zavadivian was carried out by S. Zubovich (1978;
analysis of paleowater temperature was not performed). The correlation of the depth
fluctuation curve of water bodies (water level rise/fall) located at considerable dis-
tances from each other records the general trend of water level changes caused by the
Interglacial Holsteinian/Zavadivian climate oscillations. Asynchrony (displacement
of positive and negative peaks relative to each other) and differences in the ampli-
tude of water level fluctuations in paleo-water bodies of Eastern Europe are deter-
mined by the geographical zoning of the studied territory. According to the results of
paleogeographic reconstruction of continental water bodies in Ukraine and Eastern
Europe (N 48–60°) based on ostracods, the Interglacial Holsteinian/Zavadivian had
three temperate stages (temperate I, II, III) and two cold stages (cold stage I, II). In
southern water bodies (Ukraine), three stages of warming (temperate stages I, II, III)
are recorded by an increase in water temperature, and the first and third stages of
warming (temperate I, III) are also recorded by an increase in water level. The sec-
ond warming (temperate stage II) occurs during the first prolonged cold stage (cold
stage I) and is marked by a rise in water levels in northern water bodies and an in-
crease in water temperature in southern water bodies (Fig. 23). The conclusion re-
garding climate oscillations in Holsteinian/Zavadivian (moderately warm, close to
subtropical climate with short cool and warmer periods) was also made based on
other paleontological data (theriofauna, Rekovets et al., 2014; freshwater fish, Koval-
chuk & Rekovets, 2014; mollusks, Gozhik et al., 2014) and paleopedological data
(Matviishina & Doroshkevich, 2014).
Acknowledgements. The author expresses her sincere gratitude to A. V. Matoshko,
Doctor of Geological Sciences, for organizing fieldwork on geological sections of Ukrain-
ian rivers, assistance in selecting samples for microfaunal analysis, consultation on geo-
logical and stratigraphic questions during the preparation of this article; V. N. Stepan-
N. I. Dykan
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358
chuk, Doctor of Archaeological Sciences, for inviting me to field seminars and expedition
works at the Lower Paleolithic archaeological site “Medschybisch,” for assistance in se-
lecting samples, and for providing all necessary geological materials; Tetyana Sharma for
editing and consulting on the English version of the article.
REFERENCES
Boguckyj, A. B., Łanczont, M., Łacka, B., Madeyska, T. & Nawrocki, J. 2009. Quaternary
sediment sequence at Skala Podil’ska, Dniester River basin (Ukraine): Preliminary results
of multi-proxy analyses. Quaternary International, 198, 173–194. https://doi.org/10.1016/j.
quaint.2008.05.010
Bronshtein, Z. S. 1947. Freshwater ostracods (Fauna of the USSR. Crustaceans). Academy
Sciences of the USSR, Moscow-Leningrad, 2 (1), 1–339 [In Russian].
Cohen, R. V. & Gibbard, P. L. 2019. Global chronostratigraphic correlation table for the 2.7
million years, version 2019 QI-500. Quaternary International 500, 20–31. https://doi.
org/10.1016/j.quaint.2019.03.009
Cohen, K. & Gibbard, P. 2022. Global chronostratigraphical correlation table for the last 2.7 million
years v.2019 (Poster version). Mendeley Data, V 5. https://doi.org/10.1016/j.quaint.2019.03.009
Gradstein, F. M., Ogg, J. G., Schmitz, M. & Ogg, G. М. 2020. Geologic Time Scale 2020. Published
by Elsevier BV, 2, 1–1351.
Gozhik, P. F. 2006. Freshwater mollusks of the Late Cenozoic in southern Eastern Europe.
Superfamily Unionoidea. Logos, Kyiv, 1, 1–280 [In Ukrainian].
Gozhik, P. F., Bogucki, A. B., Dmytruk, R. Ya. & Tomeniuk, O. M. 2014. Paleogeographic conditions of
formation of deposits with Pleistocene malacofauna of the Medschybisch section (Khmelnytsky
region). Ed. V. N. Stepanchuk. Scientific Bulletin “Medzhybizh” — 1’2014: Medzhybizh Locality
and problems of Lower Palaeolithic studies of the East European plain. Collection of scientific
paper. Medzhybizh-Ternopol-Kyiv: CLL “Terno-Graf”, Ternopil, 2, 79–83 [In Ukrainian].
De Deckker, P. 1979. The Middle Pleistocene ostracod fauna of the West Runton freshwater begs
Norfolk. Palaeontology, 22 (2), 292–316.
Dykan, N. I. 1994 а. First information about Quaternary ostracods from the village of Hun’ky
(Middle Dnieper region). Reports of the National Academy of Sciences of the Ukraine, 6,
85–90 [In Russian].
Dykan, N. I. 1994 b. First information about Quaternary ostracods from the village of Melnyky
(Middle Dnieper region). Reports of the National Academy of Sciences of the Ukraine, 10,
83–86 [In Russian].
Dykan, N. I. 1995. Palaeogeographic conditions in the estuary of the Donau River during the
Pre-Euxine transgression. Collection of the Institute of Geology and Mineral Resources of
the National Academy of Sciences of the Ukraine “Fossil organisms of the Phanerozoic era in
Ukraine”. Manuscript, Kyiv, 161–166 [In Russian].
Dykan, N. I. 1996. Zonocypris horizon — stratigraphic marker of Middle Pleistocene deposits of the
Interglacial Likhvin in the Azov region. Abstracts of the XIX session of the UPA “Biostratigraphic
research in the search for mineral resources in Ukraine”. Institute of Geological Sciences of the
National Academy of Sciences of the Ukraine, Kyiv, 64–66 [In Russian].
Dykan, N. I. 2001 а. Freshwater ostracods from alluvial Pleistocene deposits of Ukraine. Paleonto
logical Collection, 33, 32–49 [In Ukrainian].
Dykan, N. I. 2001 b. Key section of the Interglacial Mindel-Riss in the Middle Dnieper region.
Palaeontological justification of stratigraphic units of the Phanerozoic in Ukraine, Institute
of Geological Sciences of the National Academy of Sciences of the Ukraine, Kyiv, 91–94.
Dykan, N. I. 2003. Biostratigraphic division of Neopleistocene deposits in Ukraine based on
ostracods. Geological Journal, 3, 114–120 [In Ukrainian].
Paleogeographic Reconstruction of the Continental Water Bodies of Ukraine
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 4
359
Dykan, N. I. 2005. Use of ecological analysis methods for practical stratigraphy of Quaternary
deposits. Biostratigraphic criteria for the division and correlation of Phanerozoic deposits.
Institute of Geological Sciences of the National Academy of Sciences of the Ukraine, Kyiv,
270–275 [In Ukrainian].
Dykan, N. I. 2006. Systematization of Quaternary Ostracoda of Ukraine (a reference book).
Institute of Geological Sciences of the National Academy of Sciences of the Ukraine, Kyiv,
1–430 [In Ukrainian, English].
Dykan, N. I. 2008. Neogene-Quaternary ostracods of the Carpathian Foothills. Fourth Wave,
Kyiv, 1–88 [In Ukrainian].
Dykan, N. I. 2012. The history of ostracod development and palaeogeography of the Euxine
Basin in the Middle Miocene-Early Pliocene. Geological Journal, 1, 57–66 [In Russian].
https://doi.org/10.30836/igs.1025-6814.2012.1.138247
Dykan, N. I. 2014. First data on fossil ostracods (class Crustacea, subclass Ostracoda) from
alluvial deposits of Deer excavation area of archaeological locality Medzhybizh 1. Scientific
Bulletin “Medzhybizh” – 1–2014: Medzhybizh Locality and problems of Lower Palaeolithic
studies of the East European plain. Collection of scientific paper. Medzhybizh-Ternopol-
Kyiv: CLL “Terno-Graf”, Ternopil, 2, 85–88 [In Ukrainian].
Dykan, N. I. 2015. Palaeogeographic reconstruction of the biotopes of the Southern Bug River
at the Lower Palaeolithic locality of Medsczhybizh according to ostracods. Materials of the
XIX Ukrainian-Polish seminar “Lesy and the Palaeolithic of Podillia”, Ivan Franko National
University, Lviv, 55–56 [In Ukrainian].
Dykan, N. I. 2016. Neogene-Quaternary ostracods of the northern Black Sea. Chetverta Hvilya,
Kiev, 1–272 [In Russian].
Dykan N. I. 2023. Biostratigraphy of Neogene and Quaternary deposits (Middle Miocene-
Holocene) of the Northern part of the Black Sea according to ostracods. Collection of
scientific works of the Institute of Geological Sciences NAS of Ukraine, 16 (2), 37–51.
https://doi.org/10.30836/igs.2522-9753.2023.295227 [In Ukrainian]
Dykan N. I. 2026. New data on methods for palaeogeographic reconstructions of ancient
environments using ostracods (Arthropoda, Crustacea), Zoodiversity, 60 (3), 280–291.
https://doi.org/10.15407/zoo2026.03.280
Fuhrmann, R. 2012. Atlass of Quaternary and Recent Ostracods of Middle Germany. Naturkund
liches Museum Mauritianum, Altenburg/Thüringen, Germany, 15, 1–320 [In German].
Karmishina, G. I. 1966. Microfaunal characteristics of Neogene deposits in the Orenburg Priural
region. Issues of Geology in the Southern Urals and Volga region, Saratov University Press,
Saratov, 62, 71, 272 [In Russian].
Kovalenko, A. L. 1976. Recent ostracods of the Dniester basin. Shtiintsa, Kyshynev, 1–180 [In
Russian].
Kovalenko, A. L. 1987. Bentocypria — a new genus of the superfamily Cypridacea (Crustacea,
Ostracoda). Stratigraphy of the Upper Phanerozoic of Moldova. Shtiintsa, Kyshynev, 99–105
[In Russian].
Kovalenko, A. L. 1988. Candonidae (Ostracoda) of the southwestern of the USSR. Shtiintsa,
Kyshynev, 1–175 [In Russian].
Kovalchuk, O. M. & Rekovets, L. I. 2014. Paleoecology of the fauna of the Medzhybizh locality on
the basis of studing of remains freshwater fishes. Scientific Bulletin “Medzhybizh” — 1’2014:
Medzhybizh Locality and problems of Lower Palaeolithic studies of the East European plain.
Collection of scientific paper. Medzhybizh-Ternopol-Kyiv: CLL “Terno-Graf”, Ternopil, 2,
89–92.
Krokhmal, O. I. & Rekovets L. I. 2010. Locations of small mammals of the Pleistocene in Ukraine
and adjacent territories. LAT & K, Kyiv, 1–330 [In Russian].
Krstić, N. 1985. New specia of the Quarternary ostracodes from Vojvodina. Proceedings
Geoinstitute, 18, 197–206.
N. I. Dykan
ISSN 2707-725X. Zoodiversity. 2026. Vol. 60, No. 4
360
Matoshko, A. V., Gozhik, P. F. & Ivchenko, A. S. 2002. The fluvial archive of the Middle and
Lower Dnieper (a review). Netherland Journal Geosciences, 81 (3–4), 339–355. https://doi.
org/10.1017/S0016774600022642
Matviishina, Zh. N. & Doroshkevich, S. P. 2014. Nature conditions of the territory of Pobuzhye
in Zavadovka time (Middle Pleistocene) after data of investigations of fossil soils. Scientific
Bulletin “Medzhybizh” — 1’2014: Medzhybizh Locality and problems of Lower Palaeolithic
studies of the East European plain. Collection of scientific paper. Medzhybizh-Ternopol-
Kyiv: CLL “Terno-Graf”, Ternopil, 2, 111–119 [In Ukrainian].
Matviishina, Zh. N. & Karmazinenko, S. P. 2014. Results of paleopedological studing of Quaternary
deposits of Medschybisch Paleolithic locality. Scientific Bulletin “Medzhybizh” — 1’2014:
Medzhybizh Locality and problems of Lower Palaeolithic studies of the East European plain.
Collection of scientific paper. Medzhybizh-Ternopol-Kyiv: CLL “Terno-Graf”, Ternopil, 2,
49–69 [In Ukrainian].
Meisch, C. 2000. Freshwater Ostracoda of Western and Central Europe /Süßwasserfauna von
Mitteleuropa. 8, Crustacea (1, H 3), Heidelberg, Berlin, 1–522 [In German].
Negadaev-Nikonov, K. N. 1957. Representatives of the genus Limnocythere from Pliocene
and Quaternary deposits of the Lower Kama basin. Scientific Notes. Geological Collection.
Kyshynev University Press, Kyshynev, 25, 47–52 [In Russian].
Negadaev-Nikonov, K. N. 1974. Ostracods of the continental Pleistocene of the southern
European part of the USSR. Shtiintsa, Kishinev, 1–214 [In Russian].
Reimers, N. F. 1991. Popular Biological Dictionary. Nauka, Moscow, 1–535 [In Russian].
Reimers, N. F. 1994. Ecology, theory, laws, rules, principles and hypotheses. Nauka, Moscow,
1–367 [In Russian].
Recovets, L., Chepalyga, A. & Povodyrenko, V. 2007. Geology and mammalian fauna of the
Middle Pleistocene site, Medzhybizh, Ukraine. Quaternary International, 160, 70–80.
https://doi.org/10.1016/j.quaint.2006.09.014
Rekovets, L. I., Socha, P., Stepanchuk, V., Kovalchuk, O. & Demeshkant, V. I. 2014. Reconstruction
of existence conditions of theriofauna and ancient man during the Likhvin epoch at the
Medzhybizh locality in Ukraine. Scientific Bulletin “Medzhybizh” — 1’2014: Medzhybizh Locality
and problems of Lower Palaeolithic studies of the East European plain. Collection of scientific
paper. Medzhybizh-Ternopol-Kyiv: CLL “Terno-Graf”, Ternopil, 2, 70–79 [In Ukrainian].
Schneider, G. F. & Konstantinova, I. A. 1966. Ostracods from Pleistocene deposits of the
southwestern USSR and their significance for stratigraphy. Bulletin of the Committee for the
Study of the Quaternary Period. Nauka, Moscow, 32, 30–39 [In Russian].
Stepanchuk, V. (ed.). 2014. Scientific Bulletin “Medzhybizh” — 1–2014: Medzhybizh Locality and
problems of Lower Palaeolithic studies of the East European plain. Collection of scientific
paper. Medzhybizh-Ternopol-Kyiv: CLL “Terno-Graf”, 1–2014, Ternopil, 2, 1–260 [In
Ukrainian, English, Russian].
Stepanchuk, V. N., Ryzhov, S. N., Matviishina, Zh. N. & Karmazinenko, S. P. 2012. Report on the
2011 excavations near the village of Medzhybizh, Letychivsky District, Khmelnytsky Region.
Institute of Archaeology of the Academy Sciences of theUkraine, Kyiv, 1–145.
Stepanchuk, V. N., Ryzhov, S. N., Matviishina, Zh. N., Karmazinenko, S. P. & Moigne, A. M.
2014. First results of investigation of Medzhybizh Lower Paleolithic location, 2014. Scientific
Bulletin “Medzhybizh” — 1–2014: Medzhybizh Locality and problems of Lower Palaeolithic
studies of the East European plain. Collection of scientific paper. Medzhybizh-Ternopol-
Kyiv: CLL “Terno-Graf”, Ternopil, 2, 22–49 [In Russian].
Stratigraphic Code of Ukraine. 2012. Ed. P. Hozhyk, 2-nd edition, Logos, Kyiv, 1–66.
Zubovich, S. F. 1978. Ostracods of Middle Pleistocene deposits in Belarus and southern Lithuania.
Science and Technology, Minsk, 1–167 [In Russian].
Received 22 September 20205
Accepted 12 August 2026
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Appendix
Fig. 1. Map-scheme of the study of alluvial deposits of rivers in Ukraine: 1 — Zbruch River, seventh flood-
plain terrace of the Dniester basin, Burdyakivtsi village, cross-section “Scala Podilska”, Ternopil Region,
Lower Pleistocene, Gelasian (Tiglian/Beregovian) — Calabrian (Eburonian/Berezanian) (Dykan, 2008);
2 — Lower Danube, Lake Kahul, fifth floodplain terrace of the Danube River, Nagorne village, Odesa Re-
gion, Middle Pleistocene, Chibanian (Cromerian complex, Interglacial III-IV/Lubenian) (Dykan, 1995);
3 — Lower Danube, Lake Yalpukh, fourth floodplain terrace of the Danube River, Oserne village, Ode-
sa Region, Middle Pleistocene, Chibanian (Holsteinian/Davnievksinian) (Dykan, 1995, 2001 a, 2003,
2006); 4 — Psel River, fourth floodplain terrace of the Dnipro River basin, Hun’ky village, cross-sec-
tion “Hun’ky”, Poltava Region, Middle Pleistocene, Chibanian (Holsteinian/Zavadivian) (Dykan, 1994 a,
2001, 2006); 5 — Dnipro River, fourth floodplain terrace, Melnyky village, cross-section “Melnyky”,
Cherkasy Region, Middle Pleistocene, Chibanian (Holsteinian/Zavadivian) (Dykan, 1994 b, 2001, 2006);
6 — Southern Bug River, fourth floodplain terrace, Medschybisch town, cross-section “Medschybisch”,
Khmelnytskyi Region, Middle Pleistocene, Chibanian (Holsteinian/Zavadivian) (Dykan, 2014, 2015);
7 — Molochna River, fourth floodplain terrace, Molochna village, Zaporizhzhia Region, Middle Pleisto-
cene, Chibanian (Holsteinian/Zavadivian) (Dykan, 1996, 2006); 8 — Berda River, fourth floodplain ter-
race, Ossypenko village, Zaporizhzhia Region, Middle Pleistocene, Chibanian (Holsteinian/Zavadivian)
(Dykan, 1996, 2006); 9 — Dnipro River, third floodplain terrace, Vyschenky village, Kyiv Region, Late
Pleistocene (Eemian/Kaydakian) (Dykan, 2006); 10 — Dnipro River, second floodplain terrace, Stare
village, Kyiv Region, Late Pleistocene (Eemian/Prylukian) (Dykan, 2006); 11 — Dnipro River, second
floodplain terrace, Trypillja town, Kyiv Region, Late Pleistocene (Eemian/Prylukian) (Dykan, 2006);
12 — Molochna River, second floodplain terrace, Kostjantynivka village, Zaporizhzhia Region, Late
Pleistocene (Eemian/Prylukian) (Dykan, 2006); 13 — Noryn’ River, a left tributary of the Uzh River, the
first floodplain terrace of the Prypjat River basin, Sorokopen village, Zhytomyr Region, Late Pleistocene
(Weichselian/Prychernomorsk) (Dykan, 2006); 14 — Sula River, the first floodplain terrace of the Dni-
pro basin, Lubny city, Poltava Region, Late Pleistocene (Weichselian/Prychernomorsk) (Dykan, 2006);
15 — Ros River, the first floodplain terrace of the Dnipro River basin, Maslivka village, Kyiv Region,
Late Pleistocene (Weichselian/Prychernomorsk) (Dykan, 2006); 16 — Desna River, a left tributary of the
Dnipro, modern floodplain, Slabyn village, Zhytomyr Region. Holocene (Dykan, 2006); 17 — Dnipro
River, Kaniv Reservoir, modern floodplain, Lyplyave village, Cherkasy Region, Holocene (Dykan, 2006);
18 — Vorskla River, left tributary of the Dnipro, modern floodplain, Krotenky (Semеnivka) village, Pol-
tava Region, Holocene (Dykan, 2006); 19 — Prut River, modern floodplain, Kolomyia, Ivano-Frankivsk
Region, Holocene (Dykan, 2006)
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Fig. 2. Geological section “Scala Podilska” (Profiles A, B): I — modern landslide in the central
part of the quarry; II — location of the main profile A (Zbruch River, Western Ukraine) (photo
courtesy of A. Bogutskyi)
II
I
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Fig. 3. Geological structure of the Paleozoic-Quaternary deposits of the “Scala Podilska” section
(Profile A). Conventional notation: 1 — gravel deposits of basal alluvium; 2 — pebbles, gravel;
3 — modern soil; 4 — paleontological remains; 5 — paleosoil; 6 — loam, 7 — limestone, 8 —
sandstones, 9 — sand, 10 — clay, 11 — loess; 12 — moraine; 13 — silt; 14 — loamy sand
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Fig. 4. Species composition of ostracods from alluvial deposits of the “Scala Podilska” section
(Lower Pleistocene, Tiglian/Beregovian, Eburonian/Berezanian)
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Fi
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Fig. 7. Geological structure of the Archean–Quaternary deposits of the “Medschybisch” section
(Stepanchuk et al., 2012; Matviishіna & Karmazіnenko, 2014): A — Medschybisch “A” profile,
right bank of the Southern Bug River, 1 km north of the Medschybisch 1 “Reindeer” profile;
B — Medschybisch 1 “Reindeer” profile, left bank of the Southern Bug River; C — stratigraphic
profile of the “Medschybisch” section (Recovets et al., 2007). Conventional notation: 1 — gravel
deposits of basal alluvium; 2 — pebbles, gravel; 3 — modern soil; 4 — paleontological remains;
5 — paleosoil; 6 — loam, 7 — limestone, 8 — sandstones, 9 — sand, 10 — clay, 11 — loess; 12 —
moraine; 13 — silt; 14 — loamy sand
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Fig. 8. Species composition of ostracods from alluvial deposits of the “Medschybisch” section
(Middle Pleistocene, Cromerian/Martonoshian–Saalian/Dnieperian)
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Fig. 9. Reconstruction of water depth and water temperature in paleobiotopes of the Southern
Bug River according to the quantitative modification method (“Medschybisch” section, Western
Ukraine, Middle Pleistocene, Cromerian/Martonoshian–Saalian/Dnieperian). Conventional no-
tation: 1 — a species’ tolerance range to an abiotic factor; 2 — optimal values of an abiotic factors
for the existence of a species; 3 — negative values of an abiotic factors; 4 — reconstructed values
of an abiotic factors; 5 — trend of an abiotic factor; 6 — undefined range of species tolerance to
an abiotic factors; 7 — conditionally reconstructed value of an abiotic factor
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Fig. 10. Graph showing fluctuations in water depth and water temperature in the floodplain of the
Southern Bug River (“Medschybisch A” profile, Middle Pleistocene, Interglacial I–II, Cromerian
complex/Martonoshian–Saalian/Dnieperian). Conventional notation: 1 — water depth, m; 2 —
water temperature, °C
Fig. 11. Geological section of alluvial deposits of the Medschybisch-1 “Reindeer profile” (Stepan-
chuk et al., 2012; Matviishіna & Karmazіnenko, 2014; Dykan, 2014)
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Fig. 12. Taphonomy of fossil mollusk shells Unio spp. of autochthonous burial (subhorizontal and
horizontal orientation of shells and valves) at the bottom of a floodplain lake (Medschybisch-1
“Reindeer profile”, Southern Bug River, Middle Pleistocene, Holsteinian/Zavadivian) (Gozhik et
al., 2014)
Fig. 13. Graph of the fluctuations in water depth and water temperature in the floodplain of the
Southern Bug River (Medschybisch-1 ”Reindeer profile”, Middle Pleistocene, Holsteinian/Zavad-
ivian–Saalian/Dnieperian)
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Fig. 14. Geological structure of Middle-Upper Pleistocene deposits of the “Hun’ky” section, right
bank of the Psel River (left tributary of the Dnipro), Poltava Region, Central Ukraine (Dykan,
1994 a). Conventional notation: 1 — gravel; deposits of basal alluvium; 2 — pebbles, gravel; 3 —
modern soil; 4 — paleontological remains; 5 — paleosoil; 6 — loam; 7 — limestone; 8 — sand-
stones; 9 — sand; 10 — clay; 11 — loess; 12 — moraine; 13 — silt; 14 — loamy sand
Fig. 16. Species composition of ostracods from alluvial deposits of the “Hun’ky” section (Middle
Pleistocene, Chebanian, Holsteinian/Zavadivian)
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Be
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C
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Fig. 18. Graph of fluctuations in water depth and water temperature in the floodplain of the Psel
River (“Hun’ky” section, Central Ukraine, Middle Pleistocene, Holsteinian/Zavadivian). Con-
ventional notation: 1 — water depth, m; 2 — water temperature, °C
Fig. 19. Geological structure of Middle Pleistocene deposits of the ”Melnyky” section, Dnipro
River, Tscherkassy region, Central Ukraine (Dykan, 1994 b). Conventional notation: 1 — gravel
deposits of basal alluvium; 2 — pebbles, gravel; 3 — modern soil; 4 — paleontological remains;
5 — paleosoil; 6 — loam, 7 — limestone, 8 — sandstones, 9 — sand, 10 — clay, 11 — loess; 12 —
moraine; 13 — silt; 14 — loamy sand
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Fig. 20. Species composition of ostracods in alluvial and lacustrine deposits of the “Melnyky”
section (Middle Pleistocene, Chebanian, Holsteinian/Zavadivian)
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Fig. 21. Reconstruction of water depth and water temperature in paleobiotopes of the Dnipro
River according to the quantitative modification method (“Melnyky” section, Central Ukraine,
Middle Pleistocene, Holsteinian/Zavadivian). Conventional notation: 1 — a species’ tolerance
range to an abiotic factor; 2 — optimal values of an abiotic factors for the existence of a species;
3 — negative values of an abiotic factors; 4 — reconstructed values of an abiotic factors; 5 — trend
of an abiotic factor; 6 — undefined range of species tolerance to an abiotic factors; 7 — condition-
ally reconstructed value of an abiotic factor
Fig. 22. Graph of fluctuations in water depth and water temperature in the floodplain of the Dni-
pro River (“Melnyky” section, Central Ukraine, Middle Pleistocene, Holsteinian/Zavadivian).
Conventional notation: 1 — water depth, m; 2 — water temperature, °C
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Fig. 23. Correlation of bottom depth and water temperature, reconstructed according to ostra-
cods in continental water bodies of Ukraine (Dikan, 1994 a, b; 2014) and Belarus, Lithuania, and
Russia (Zubovich, 1978) in the Interglacial Holsteinian/Zavadivian
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Paleontological table 3. Freshwater ostracods from alluvial deposits of river terraces of Ukraine:
1 — Limnocythere inopinata (Baird, 1843), adult, left valve, }, external, SEM, × 220, Central
Ukraine, Psel River, “Hun’ky” section, floodplain facies, layer 5, Middle Pleistocene, Holstein-
ian/Zavadivian; 2 — Limnocythere inopinata (Baird, 1843), adult, right valve, {, external, SEM,
× 220, ibid; 3 — Cypria candonaeformis (Schweyer, 1949), adult, left valve, external, SEM, × 160,
Western Ukraine, Zbruch River, «Scala Podilska» section (profile “A”), facies of the shallow chanel
zone, layer 5, Lower Pleistocene, Calalbrian, Eburonian/Berezanian; 4 — Cyclocypris ovum (Ju-
rine, 1820), adult, right valve, external, SEM, × 150, Western Ukraine, Southern Bug River, “Med-
schybisch” section, profile «A», flood-plain facies, layer 15a, Middle Pleistocene, Holsteinian/
Zavadivian; 5 — Cyclocypris laevis (O. Müller, 1785), adult, right valve, external, SEM, × 300,
Central Ukraine, Desna River, “Slabin” section, facies of the shallow chanel zone, Holocene, mod-
ern deposits; 6 — Cyclocypris globosa (G. Sars, 1862), adult, left valve, external, SEM, × 220,
Central Ukraine, Psel River, “Hun’ky” section, flood-plain facies, layer 5, Middle Pleistocene,
Holsteinian/Zavadivian; 7 — Cyprinotus salinus (Brady, 1862), adult, left valve, external, SEM,
× 110, Western Ukraine, Southern Bug River, “Medschybisch” section, profile “A”, floodplain fa-
cies, layer 18, Middle Pleistocene, Holsteinian/Zavadivian; 8 — Cyprinotus vassoevichi Schneider
in Bodina, 1961, adult, right valve, external, SEM, × 120, Central Ukraine, Psel River, “Hun’ky”
section, floodplain facies, layer 7, Middle Pleistocene, Holsteinian/Zavadivian; 9 — Herpetocy-
pris reptans (Baird, 1835), adult, left valve, external, SEM, × 160, Central Ukraine, Dnipro River,
“Melnyky” section, lake facies, layer 5, Middle Pleistocene, Holsteinian/Zavadivian; 10 — Physo-
cypria fadeevi Dubowsky, 1927, adult, left valve, external, SEM, × 220, Central Ukraine, Dnipro
River, “Melnyky” section, lake facies, layer 6, Middle Pleistocene, Holsteinian/Zavadivian
N. I. Dykan
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Paleontological table 4. Freshwater ostracods from alluvial deposits of river terraces in Ukraine:
1 — Ilyocypris gibba (Ramdohr, 1808), adult, left valve, external, SEM, × 120, Western Ukraine,
Zbruch River, «Scala Podilska» section (profile “B”), floodplain facies, layer 6, Lower Pleisto-
cene, Calalbrian, Eburonian/Berezanian; 2 — Ilyocypris gibba (Ramdohr, 1808), juvenile, right
valve, external, SEM, × 140, ibid; 3 — Ilyocypris bradyi Sars, 1890, adult, left valve, external, SEM,
× 100, Western Ukraine, Southern Bug River, “Medschybisch” section, profile “A”, flood-plain
facies, layer 16a, Middle Pleistocene, Holsteinian/Zavadivian; 4 — Limnocythere tuberculata Ne-
gadaev, 1957, adult, right valve, {, external, SEM, × 160, Central Ukraine, Psel River, “Hun’ky”
section, floodplain facies, layer 5, Middle Pleistocene, Holsteinian/Zavadivian; 5 — Limnocythere
tuberculata Negadaev, 1957, adult, right valve, }, external, SEM, × 180, ibid; 6 — Limnocythere
tuberculata Negadaev, 1957, adult, left valve, { external, SEM, × 180, Western Ukraine, Zbruch
River, “Skela Podilska” section, profile “A”, facies of the chanel, layer 4, Lower Pleistocene, Calal-
brian, Eburonian/Berezanian; 7 — Denticulocythere dorsotuberculata (Negadaev, 1957), adult,
left valve, {, external, SEM, × 240, Central Ukraine, Psel River, “Hun’ky” section, flood-plain
facies, layer 5, Middle Pleistocene, Holsteinian/Zavadivian; 8 — Denticulocythere dorsotubercu-
lata (Negadaev, 1957), adult, right valve, }, external, SEM, × 220, ibid; 9 — Paralimnocythere
originalis (Negadaev, 1965), adult, right valve, {, external, SEM, × 180, Central Ukraine, Dnipro
River, “Melnyky” section, lake facies, layer 6, Middle Pleistocene, Holsteinian/Zavadivian; 10 —
Paralimnocythere originalis (Negadaev, 1965), adult, left valve, }, external, SEM, × 160, ibid
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Paleontological table 5. Freshwater ostracods from alluvial deposits of river terraces in Ukraine:
1 — Candona neglecta Sars, 1887, adult, left valve, }, external, SEM, × 50, Central Ukraine, Psel
River, “Hun’ky” section, floodplain facies, layer 3, Middle Pleistocene, Holsteinian/Zavadivian;
2 — Candona (С.) neglecta Sars, 1887, adult, left valve, {, external, SEM, × 50, ibid; 3 — Candona
(С.) iliensis Mandelstam, 1963, adult, right valve, }, external, SEM, × 110, Central Ukraine, Psel
River, “Hun’ky” section, flood-plain facies, layer 4, Middle Pleistocene, Holsteinian/Zavadivi-
an; 4 — Candona(С.) iliensis Mandelstam, 1963, adult, left valve, {, external, SEM, × 110, ibid;
5 — Candona (С.) elongata (Schweyer, 1949), adult, left valve, }, external, SEM, × 100, Central
Ukraine, Psel River, “Hun’ky” section, flood-plain facies, layer 5, Middle Pleistocene, Holstein-
ian/Zavadivian; 6 — Candona (С.) elongata (Schweyer, 1949), adult, left valve, {, external, SEM,
× 100, ibid; 7 — Candona (С.) angulata G. Müller, 1900, adult, left valve, }, external, SEM, × 80,
Central Ukraine, Psel River, “Hun’ky” section, flood-plain facies, layer 4, Middle Pleistocene,
Holsteinian/Zavadivian; 8 — Candona (С.) candida (O.Müller, 1776), adult, left valve, external,
SEM, × 100, Western Ukraine, Southern Bug River, “Medschybisch”section. profile “A”, floodplain
facies, layer 15 b, Middle Pleistocene, Holsteinian/Zavadivian; 9 — Candona (E.) balatonica Da-
day, 1894, adult, right valve, external, SEM, × 100, Central Ukraine, Dnipro River, “Melnyky”
section, lake facies, layer 5, Middle Pleistocene, Holsteinian/Zavadivian; 10 — Darwinula steven-
soni (Brady and Robertson, 1870), adult, left valve, external, SEM, × 160, Central Ukraine, Psel
River, “Hun’ky”“ section, flood-plain facies, layer 5, Middle Pleistocene, Holsteinian/Zavadivian
N. I. Dykan
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Paleontological table 6. Freshwater ostracods from alluvial deposits of river terraces in Ukraine:
1 — Typhlocypris compressa (Kosh, 1837), adult, right valve, {, external, SEM, × 130, Central
Ukraine, Dnipro River, “Melnyky” section, lake facies, layer 7, Middle Pleistocene, Holsteinian/
Zavadivian; 2 — Typhlocypris compressa (Kosh, 1837), adult, left valve, }, external, SEM, × 160,
ibid. 3 — Typhlocypris rostrata (Brady and Norman, 1889), adult, left valve, external, SEM, × 120,
Western Ukraine, Southern Bug River, “Medschybisch” section, profile «A», flood-plain facies,
layer 15, Middle Pleistocene, Holsteinian/Zavadivian. Species-indicators of the Sarmatian: 4 —
Callistocythere molesta Bonaduce, 1988, adult, left valve, external, SEM, × 100, Western Ukraine,
Zbruch River, “Skala Podilska” section, profile “A”, marine deposits, layer 8, Middle Miocene,
Serravallian/Sarmatian; 5 — Aurila sarmatica (Zalányi, 1949), adult, left valve, }, external, SEM,
× 100, ibid; 6 — Aurila sarmatica (Zalányi, 1949) adult, right valve, male, external, SEM, х 100,
ibid; 7 — Hemicytheria omphalodes (Reuss, 1854), adult, left valve, }, external, SEM,100, ibid;
8 — Hemicytheria omphalodes (Reuss, 1854) adult, left valve, {, external, SEM, × 100, ibid; 9 —
Haplocytheridea dacica (Hejas, 1894) adult, right valve, } external, SEM, × 100, ibid.; 10 — Loxo-
concha ornata Schneider, 1939, adult, right valve, external, SEM, × 100, ibid
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| id | oai:ojs.akademperiodyka.org.ua:article-1042 |
| institution | Zoodiversity |
| issn | 2707-7268 |
| keywords_txt_mv | |
| language | English |
| last_indexed | 2026-08-27T01:00:40Z |
| publishDate | 2026 |
| publisher | Publishing House "Akademperiodyka" of the National Academy of Sciences of Ukraine |
| record_format | ojs |
| resource_txt_mv | ojsakademperiodykaorgua/56/41e22417211d75de8786e3e4c253aa56.pdf |
| spelling | oai:ojs.akademperiodyka.org.ua:article-10422026-08-26T09:51:17Z Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) Dykan , N. I. freshwater ostracods ecology Eastern Europe paleoclimate Holsteinian A paleogeographic reconstruction of Quaternary continental water bodies in Ukraine based on ostracods requires the data necessary for correlating the stages of development of water bodies in Eastern and Western Europe to be generalised, and synchronised with the general climatic trend of the Quaternary. A variety of methods and analyses were employed, including actualistic, quantitative modification, taphonomic, statistical, population, ecological, zoogeographical and biostratigraphic approaches. The detailed reconstruction of river and lake paleo-biotopes (type of biotope, degree of trophicity, water depth of biotope, m; water temperature , °C) in the paleo-valleys of the Zbruch, Southern Bug, Psol and Dnipro rivers during the Early to Middle Pleistocene (Calabrian – Holsteinian) has been conducted. A comparative analysis of the development of continental water bodies in Ukraine, Lithuania, Belarus and Central Russia during the Middle Pleistocene (Holsteinian) was conducted based on the results of studying freshwater ostracods of Eastern Europe. Correlating fluctuations (increases/decreases) in the water levels of paleo-water bodies located at considerable distances from each other reveals a general trend in water level changes caused by Interglacial Holsteinian climatic oscillations. It also reveals asynchrony (the partial displacement of positive and negative peaks relative to each other) and different water level fluctuation amplitudes caused by geographical zoning of the territory. According to the ostracods, the Holsteinian interglacial period had three warm stages (temperate stages I, II and III) and two cold stages (cold stages I and II). In southern water bodies (Ukraine), three stages of warming (temperate stages I, II and III) are recorded by an increase in water temperature. The first and third stages of warming (temperate stages I and III) are also recorded by an increase in water level. The second period of warming (temperate stage II) occurred during the first prolonged cold stage (cold stage I), which is indicated by rising water levels in northern bodies of water and only by an increase in water temperature in southern water bodies. Based on ostracods the timing of the formation and dynamics of a paleo-landslide in the paleo-valley of the Zbruch River during the Early Pleistocene (Calabrian) have been reconstructed. Publishing House "Akademperiodyka" of the National Academy of Sciences of Ukraine 2026-08-07 Article Article application/pdf https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/1042 10.15407/zoo2026.04.342 Zoodiversity; Vol. 60 No. 4 (2026): Zoodiversity Zoodiversity (Vestnik Zoologii); Том 60 № 4 (2026): Zoodiversity 2707-7268 2707-725X 10.15407/zoo2026.04 en https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/1042/407 Copyright (c) 2026 N. I. Dykan |
| spellingShingle | Dykan , N. I. Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) |
| title | Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) |
| title_full | Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) |
| title_fullStr | Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) |
| title_full_unstemmed | Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) |
| title_short | Paleogeographic reconstruction of the continental water bodies of Ukraine in the Early-Middle Pleistocene based on ostracods (Arthropoda, Crustacea) |
| title_sort | paleogeographic reconstruction of the continental water bodies of ukraine in the early-middle pleistocene based on ostracods (arthropoda, crustacea) |
| topic_facet | freshwater ostracods ecology Eastern Europe paleoclimate Holsteinian |
| url | https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/1042 |
| work_keys_str_mv | AT dykanni paleogeographicreconstructionofthecontinentalwaterbodiesofukraineintheearlymiddlepleistocenebasedonostracodsarthropodacrustacea |