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Article

Paleoecological Reconstruction Derived from an Age–Depth Model and Mollusc Data, Pécel, Hungary

by
László Makó
1,*,
Péter Cseh
2,
Balázs Nagy
2,
Pál Sümegi
2 and
Dávid Molnár
2
1
Institute of Plant Sciences and Environmental Protection, Faculty of Agriculture, University of Szeged, Andrássy Street 15, H-6800 Hódmezővásárhely, Hungary
2
Department of Geology, Faculty of Science and Informatics, University of Szeged, Egyetem Street 2-6, H-6722 Szeged, Hungary
*
Author to whom correspondence should be addressed.
Quaternary 2025, 8(3), 37; https://doi.org/10.3390/quat8030037
Submission received: 12 May 2025 / Revised: 24 June 2025 / Accepted: 4 July 2025 / Published: 9 July 2025
Browse Figures
Versions Notes

Abstract

The Pécel loess–paleosol profile is a 25.72-metre-high well-preserved sequence in the northern part of Hungary. It was sampled every 4 cm for the purpose of sedimentological analysis and every 12 cm for the purpose of mollusc investigation, which are relatively high resolutions in loess investigation. Twenty samples were radiocarbon-dated from the L1 layer (top 8 m of the sequence). Subsequently, an age–depth model was constructed, from which an accumulation rate was calculated. Based on these radiocarbon and previous magnetic susceptibility data, the Pécel’s L1 layer is correlated with the Chinese Loess Plateau’s L1 layer and the MIS 2–4 stages. The malacological examinations show that the temperature was basically warm during the development, and there was open vegetation except on the S2, S1 and L1S1 paleosol layers, where significant forest expansion was shown. With the magnetic susceptibility and the malacological data, it is possible to track the changes in the conditions through the Chinese Loess Plateau’s timeline.

1. Introduction

The analysis of loess–paleosol sequences has an essential role in the research of the Quaternary Period (e.g., [1,2,3,4,5,6,7,8,9,10]). Previous research [11,12], which included grain size composition, geochemical studies, and magnetic susceptibility measurements, showed the process of formation and relative age of the section. However, both the study of the mollusc fauna and the radiometric age data and age–depth models derived from them are essential for accurate high-resolution paleoecological studies [13,14,15,16,17].
The Pécel loess–paleosol section is located in the Carpathian Basin in Hungary (Figure 1), in the Gödöllő hills [18] at 47°29.797′ N and 19°21.235′ E (WGS 84), a foothill-positioned, open to the south section (e.g., [19,20,21,22,23]), which was sampled in 2019 and 2021 [11,12] using 10 subdivisions.
The present manuscript has been prepared for the purpose of verifying the correlation with the Chinese Loess Plateau, as assumed from the magnetic susceptibility data, with the complete malacological data set. The malacological data series can serve as a basis for future comparisons of the evolutionary history of both the Hungarian and the Eastern European sections and for the investigation of possible microenvironmental influences.

2. Materials and Methods

The 25.72 m high loess–paleosol wall was sampled at an interval of 4 cm for the laboratory measurements (LOI—Organic Carbon, Carbonate content; Grain Size Analysis and Magnetic Susceptibility) and with an interval of 12 cm (5-5 kg) for obtaining a sufficient amount of Mollusca material. The extraction method of the mollusc content from the sediment was by wet sieving followed by H2O2 preparation to remove organic matter [24]. Subsequently, the extracted material was sorted and identified; moreover, a part of it from certain depths was selected for radiocarbon measurements—each from the investigated layers and from the boundaries of them.

2.1. Absolute Dating

Twenty radiocarbon measurements were carried out in the Isotoptech laboratory in Debrecen [25,26] by using the extracted snail shells. The obtained radiocarbon ages were calibrated with the IntCal20 calibration curve [27] using the Calib 8.2 software [28].

2.2. Age–Depth Model

From the 20 available uncalibrated ages, an age–depth model was carried out with Bacon software [29]. Although age data were also derived from the top layer, S1, as it is the recent soil level, the model boundaries were set between 0.44 and 8.44 m, representing the entire L1 (L1L1, L1S1 and L1L2) section of the profile.

2.3. Accumulation Rate, Mass Accumulation Rate

The Accumulation Rate (AR) specifies the possible minimum, mean and maximum ages for a given depth, depending on its dimension (m×a−1). From the AR value, the Mass Accumulation Rate (MAR—[8]) can be calculated, which is a value multiplied by the density of the dry powder material (1.5—[8]) and the mass concentration of the loess (1—[8,30]), with a dimension of g×m−2×a−1. The density of the soil is different from that of the loess, and since we concentrated on the loess formation, we only made the calculations for continuity, but did not take it into account.

2.4. Magnetic Susceptibility Measurement

The magnetic susceptibility (MS) measurement (e.g., refs. [31,32,33,34,35,36,37,38] was carried out using a Bartington MS2 K instrument [13]. Every sample from 4 centimetres was dried out, powdered, and then measured in different directions three times.

2.5. Malacological Examination Method

The malacological sampling, cleaning, and identification were carried out according to the methods by Endre Krolopp and Pál Sümegi [24,39,40]. At the Pécel brickyard, ca. 5 kg of bulk samples were collected from every 12 cm intervals of the sequence from 0 to 2280 cm, resulting in 190 sediment samples. Then, the sediment was washed through sieves with a mesh size of 0.8 mm, dried, and the identifiable shells and fragments were sorted and identified. Identification and the knowledge of each species’ climatic and environmental demands can help create paleoecological reconstruction datasets [2,37,41,42,43]. This modern knowledge is used to interpret paleoenvironmental changes from fossil data, i.e., absolute and relative abundance diagrams of species and ecological groups, by applying Lyell’s Actualism [38,44,45,46]. Based on modern samplings, the ecological requirement of each species is known, and species have been classified in ecological groups that may differ from one region to another, and which are based on species temperature, humidity and vegetation preferences [2,33,39,40,47,48,49]. All three factors depend upon each other, and even the changes in other (less important) conditions.
Beyond the dominance and abundance analyses, more sophisticated statistical analyses were carried out. Any community, including the malacofauna, can be well characterised by its diversity per sample, using the Shannon index H′ [50], which is calculated as follows:
H ′ = − ∑ 1 n p k × l n   ×   p k
where n equals the number of species and pk the frequency of the k species (k varying between 1 and n).
Principal component analysis was applied to the malacological data table [51] with assemblages in the columns and species ratio per sample values in the rows. Calculated loadings for the 1st principal component provide an estimate of temperature variation, and loadings for the 2nd principal component are used to estimate the wet/dry component [52].
For the diversity counts and the multivariate statistical analyses, the Past 4.16 software kit [53] was used.

3. Results

The description and analysis of the LOI and Grain Size Analysis have been published in two previous articles [11,12], so the results that are relevant for the present publication are described. For the nomenclature separation of the loess–paleosol layers, the Chinese Loess Plateau was used (e.g., [54,55,56,57,58,59]).

3.1. Absolute Dating, AR, MAR

The model was created with an interval of 12 cm (according to the sampling) and two hiatus layers at 300 and 564 cm (according to the layer boundaries), based on 13.875 million iterations from 72 sections (Figure 2), from which the accumulation values per centimeter, the ages and the accumulation rate were plotted; 79% of the data provided fall within the 95% confidence interval.
Table 1 shows the 20 measured age data and the investigated species from which they were measured. The first of these unused data originated from the recent soil; moreover, 1-1 from the paleosol and the lower loess body. In these cases, the values of these outliers can be considered as measurement anomalies or the pollution of the samples.
The accumulation rate (Figure 3) for L1L2 (8.84–5.64 m) indicates a slow accumulation of 0.3–0.5 mm/year. In the case of the L1S1 (5.64–3.00 m) paleosol layer above it, these values are between 1.5 and 2.0 mm/year; however, soil development is the predominant process against the dust fall activity. Higher (around 1 mm/year) accumulation rates can be observed in the lower parts of the L1L1 (3.00–0.44 m) loess body, while it decreases to 0.2–0.3 mm/year towards the recent soil surface.
Since the MAR value aims to determine the falling dust, the section of the paleosol layer is covered with a grey rectangle. The average value of MAR within the L1L2 loess body (8.84–5.64 m) is 459, while in the L1L1 (3.00–0.44 m), it can be separated into two subsections: the lower (3.00–1.41 m) is 1592—this subsection is still affected by the soil development—and the upper subsection’s (1.41–0.44 m) average is 370 g×m−2×year−1. The latter average was increased by high values near the boundary of the loess and paleosol layers.

3.2. Magnetic Susceptibility

The MS curve (Figure 4—[12]) shows that the L1L2 (8.84–5.64 m) can be characterised by low values throughout the entire loess body, with an average value of 31. Subsequently, generally low values can be observed also in the case of the L1S1 (5.64–3.00 m) paleosol layer compared to the average paleosols. It can be characterised by higher and lower fluctuations; however, the average values do not exceed 38. The L1L1 (3.00–0.44 m) loess body shows continuously increasing values in the direction of the recent surface soil, with an average value of 36.
The MS curve of Pécel compared to the Chinese Loess Plateau’s MS curve can give information about the relative ages in the lower sections, which can be traced till the L3 loess section. Underlain, thin loess (L4) and paleosol (S3, S4) layers can be traced. However, unusually for loess, high MS values are found in the upper part of the L5 layer.

3.3. Malacological Examinations

The malacological studies of the Pécel loess–paleosol section do not cover the whole section, only up to 0–22.80 m. In this section, 190 samples were collected with a 12 cm sample interval according to the fine stratigraphic sampling procedures, from which 5777 statistically valuable snail shells were identified. The average number of individuals per sample is 30.4, which is considered low. Among the samples, 30 contained more than 50 shells, and 16 samples contained more than 100 shells. The highest abundance was 638 shells, between 552 and 564 cm. The shells belonged to 28 species of terrestrial snails. The average number of species per sample was 3.3, while the two most abundant samples contained 16 species, between ca. 5.50 and 5.80 m (Figure 5).

3.4. Malaco-Zones

For the malacological examination, 10 malaco-zones (PCMZ—PéCel Malaco-Zone) were assigned in the section based on the percentage distribution of snail species per sample (Figure 6). Where possible, the zones were also assigned absolute ages based on the age–depth model produced (Figure 2).
  • PCMZ-1 (22.80–21.72 m): The lowest and presumably oldest zone is located in the PL5 loess layer. The gastropod fauna is characterised by an absolute dominance of the mesophilous species (~75%; Figure 5), the assemblage being mainly composed of Vallonia costata shells. In addition, V. pulchella and the warmth-loving Helicopsis striata and Pupilla triplicata represented a more significant proportion of the fauna. Cold-resistant species were only present in an indicative proportion (Pupilla sterrii ~1%), and cold-loving species were absent. The mean abundance of the zone was 41.5, higher than the overall mean abundance, with species richness per sample ranging from one to six.
  • PCMZ-2 (21.72–18.00 m): The second malacological zone includes the upper part of the PL5 loess layer, the whole of PS4, PL4 paleosol and loess layers, and the lower part of the PS3 paleosol layer. The zone contains an approximately equal distribution of thermophilous (H. striata, P. triplicata, Chondrula tridens) and wide-tolerant species (V. costata, Pupilla muscorum); cold-resistant and cold-loving species were not found in this zone. The average abundance of the malaco-zone is low, only eight, while the frequency of species varied between one and five.
  • PCMZ-3 (18.00–15.72 m): The third zone covers the upper part of the PS3 paleosol layer, the PL3 loess layer, and the smaller lower part of the PS2 paleosol. The upper part of the zone was found to be sterile, while the rest of the zone was characterised by a high distribution of thermophilous species, especially H. striata. In addition to H. striata, P. triplicata and the mesophilous P. muscorum and V. costata were more abundant, and the cold-resistant Succinella oblonga was also present in two samples. Cryophilous species were not present in the zone. The distribution of the richness of species per sample and the mean abundance values are similar to those of PCMZ-1 (1–7 and 41.3, respectively).
  • PCMZ-4 (15.72–12.96 m): The zone is located in the upper part of the PS2 paleosol and the lower part of the PL2 loess layer. In the lower part, there is a rather high proportion of mesophilous species (Limacidae, V. pulchella, P. muscorum), and then, moving upwards, of more thermophilous species (H. striata, P. triplicata). In the lower part of the zone, the proportion of Limacidae is more pronounced. In one sample, the cold-loving Columella columella also appears. The average abundance of the zone, thanks also to the 10 sterile samples, is only 8.2, with species numbers ranging from zero to eight.
  • PCMZ-5 (12.96–9.48 m): The fifth zone covers the upper part of the PL2 layer and the upper part of the PS1 paleosol. The zone is characterised by low abundance, with an average of only one specimen per sample and species richness ranging from zero to three. Fifteen samples in the zone did not contain any valuable snail shells. Despite the low values, it can be concluded that warmth-loving species were also the most abundant in the zone, with Granaria frumentum being the most abundant species. In addition, the shells of H. striata, P. triplicata and C. tridens were also found. P. muscorum and V. pulchella are subordinately represented in the wide-tolerant species, and shells of Clausilia dubia and Nesovitrea hammonis also appear for the first time in this zone. Among the cold-resistant species, S. oblonga and Discus ruderatus, which also appear here for the first time, are present, with no cryophilous element in the zone.
  • PCMZ-6 (9.48–6.00 m; 49,172 (from 8.84 m) −33,235 cal BP): In the next zone, which includes the upper part of the PS1 layer and most of the PL1L2 loess layer, the most abundant species are the widely tolerant ones, with a high proportion of V. costata, P. muscorum and Limacidae fossils. The thermophilous species are represented, to a lesser extent, by the pair H. striata and P. triplicata, and of the cold-resistant species, only S. oblonga is present. The mean abundance is only 23.4, the species number varies from zero to five, and only one sample in the zone was sterile. Based on the age–depth model, the 884–600 cm section of the zone covers the interval 49,172–33,235 cal BP.
  • PCMZ-7 (6.00–4.68 m; 33,235–25,359 cal BP): This small (only 132 cm) zone covers only 7876 cal BP ys, and includes the upper part of PL1L1 and the lower part of the PL1S1 paleosol layer. In general, it has the highest average abundance in the section (170.3) and is also the most species-rich (5–16 species) zone on average. The malacofauna is dominated by species with wide tolerances (75–80%), with the most significant occurrence of P. muscorum and V. costata, but other species of wide tolerance are also present in this zone, which have not been found so far (Clausilia pumila, Euconulus fulvus) or only sporadically (Vallonia pulchella, Clausilia dubia, Nesovitrea hammonis, Limacidae). The warmth-loving snail fauna, similar to PCMZ-1, comprised 15–20% of the snail community, with Cochlicopa lubricella and Fruticicola fruticum present along with H. striata and P. triplicata. The proportion and number of species of cold-resistant (Trochulus hispidus, S. oblonga and Pupilla sterrii) and cold-loving (Columella columella) snails are also noteworthy.
  • PCMZ-8 (4.68–2.40 m; 25,359–23,796 cal BP): The zone covers the upper part of the PL1S1 paleosol layer and the lower part of the PL1L1 loess layer, covering about 1563 cal BP years according to the age–depth model. The average abundance of the zone is about twice the average for the whole section, 64.2, with species richness ranging from 2 to 10. The malacofauna of the zone is characterised by a significant thermophilous dominance, ranging from 50 to 90%. The highest proportions of H. striata and P. triplicata were found, but also G. frumentum, C. tridens and C. lubricella are remarkable in some samples. Mesophilous species are subordinate in the zone with a proportion of 20–50%, with the most significant presence of P. muscorum. Also remarkable are the proportions of Vallonia pulchella, C. dubia and Limacidae, and Punctum pygmaeum is also present in this zone. V. costata shells were not found in this zone. Among the cold-resistant species, Vestia turgida is the only one to occur here, and the cold-loving species are represented by two species (C. columella and Vertigo pygmaea, which appear here), but in low proportions.
  • PCMZ-9 (2.40–1.20 m; 23,796–22,235 cal BP): The zone, which is only 120 cm thick and is exclusively assigned to the PL1L1 loess layer, covers a period of 1561 cal BP years according to the age–depth model. The abundance per sample here is only 13.3 specimens, and the species number varies from one to five. This zone is also characterised by the dominance of warmth-loving species (30–100%), with only H. striata and P. muscorum present, the former in several samples with proportions of around 50% and even 100%. Mesophilous species are represented in lower proportions but with more species. In addition to P. muscorum, V. costata reappears, with a predominant proportion of this pair. In addition, C. dubia, E. fulvus and Limacidae shells were also found in the zone. The zone contained neither cold-tolerant nor cold-loving species.
  • PCMZ-10 (1.20–0 m; 22,235–18,090 (from 44 cm) cal BP): The uppermost and youngest malaco-zone of the section covers the upper third of the PL1L1 loess layer and the PS0 modern soil layer. The abundance per sample has increased slightly (25.4), as has the species richness (2–12) compared to PCMZ-2. As we move upwards in the zone, we observe gradually increasing warmth-loving (from 40% to 80%) and gradually decreasing mesophilous (from 75% to 30%) proportions, along with more significant cold-resistant proportions (20–25%). Among the thermophilous species, the proportion of H. striata has decreased; the most dominant thermophilous species are P. triplicata and G. frumentum (mainly in the upper part of the zone), and the presence of F. fruticum is also noticeable in the zone. Among the wide-tolerant species, in addition to V. costata and P. muscorum, Vitrea crystallina and Vitrina pellucida are also present in the upper part of the zone (PS0). Three species, P. sterrii, S. oblonga and T. hispidus, make up the larger proportion of cold-tolerant species. No cryophilous species were found in this zone.

3.5. Diversity and PCA

Not surprisingly, the diversity curve of the malacofauna shows the highest similarity with the species richness curve (Figure 5). The diversity values per sample range from 0 to 2. Highest values were found in the malaco-zones PCMZ-2, PCMZ-4, PCMZ-7 and PCMZ-9.
The per-sample loadings of the first and second principal components of the principal component analysis provided estimates of temperature and ambient humidity. The first principal component loadings represent the temperature trend, and the second principal component loadings represent the humidity trend. The reconstructed temperature in the section could be essentially higher because the loading values are predominantly in the positive range. The highest temperature peaks occurred in the malaco-zones PCMZ-3, PCMZ-4, PCMZ-5 and PCMZ-9, and the lowest temperatures were reconstructed in the zones PCMZ-2, PCMZ-4 and PCMZ-5. The humidity values were not very different from zero, with drier periods in zones PCMZ-3, PCMZ-4, PCMZ-8 and PCMZ-9 and wetter periods in malaco-zones PCMZ-1, PCMZ-2, PCMZ-6 and PCMZ-9.

4. Discussion

4.1. Absolute Dating, AR, MAR

The AR values are changing between 0.25 and 0.5 mm/year in the L1L2 layer, until the soil L1S1 layer; also, the MAR values are between 100 and 700 g×m−2×a−1. The soil development processes should be treated separately from dust fall processes. MAR results are not taken into account here, as their values provide information about the mass accumulation of falling dust.
AR (0.15–3.33 mm/year) and MAR (300–1800 g×m−2×year−1) values show great variability in the L1 section, which can be divided into two subsections. The lower part of the L1 section’s (1.41–3.00 m) AR value is between 0.83 and 3.33 mm/year, and the MAR value is 1250–5000 g×m−2×a−1. This big variability can be explained by the influence of soil development. The upper part of the L1 section (0.44–1.41 m) shows lower values, as the AR is between 0.15 and 0.48 g×m−2×a−1. The MAR values lie between 224 and 714 g×m−2×a−1. This subsection is not affected by the soil development; thus, in the upper part, the accumulation rate continuously decreases till the boundary of the recent soil layer (S0).

4.2. Malacology

The malacological analysis of the Pécel loess–paleosol section was carried out on 190 samples collected between 0 and 22.80 m. The abundance, dominance and species richness per sample were determined. From these data, statistical analyses were performed, which included diversity and principal component analysis studies. The total number of individuals (5777 specimens) and the average number of specimens per sample (30.4 individuals per sample) are well below the numbers of specimens from much smaller sections from other areas of the Carpathian Basin [13,14,20,60,61,62,63]. However, the malacological analysis can still provide a good estimate of the development of the paleoclimate and paleoenvironment. The uniqueness of the Pécel loess–paleosol section from the malacological point of view lies in the fact that no other section of this size has been processed in such detail in the wider environment of the site.
Based on the percentage of species distribution (dominance) values per sample, 10 malaco-zones (PCMZ) were delineated in the section (Figure 5 and Figure 6). The malaco-zones also have temperature and humidity estimates, species richness, number of individuals per sample and diversity values. The calculated age–depth model also provides the absolute age of the malaco-zones in the upper, younger part of the section (8.84–0.44 m). The age of the zones in other parts of the section can only be estimated by stratigraphic correlation.
  • The snail fauna of PCMZ-1 (22.80–21.72 m) is predominantly composed of the mesophilous species Vallonia costata and Vallonia pulchella shells. V. costata tolerates moisture, V. pulchella requires moisture, so the subordinate proportion of drought-tolerant species and the absence of shade-loving species suggest that this zone represents an open, higher humidity, temperate climate. This is confirmed by the climate and humidity data as well. The zone covers the middle to lower part of the PL5 loess layer, and the correlation suggests an age of more than 400 kys, which is consistent with MIS 12 according to Marine Isotope Stages [64,65].
  • The changes in the malacofauna of PCMZ-2 (21.72–18.00 m) (high proportion of thermophilous species) indicate a warming period. In addition, the presence of a high proportion of Helicopsis striata, Pupilla triplicata and Chondrula tridens, which prefer dry, open vegetation, and of the mesophilous V. costata and Pupilla muscorum, which prefer open vegetation, indicates a dry steppe environment, which, based on the remains of the shade-loving Limacidae, which appear at two levels, may have been interspersed periodically with small tree communities. This is also supported by the graphs indicating higher temperatures and lower humidity. The malaco-zone covers the upper part of PL5, the PS4, PL4 layers completely, and the PS3 paleosol layer almost completely, so the correlation suggests an age of approximately 400–280 kys. The period of formation is thus estimated to be between MIS 12-9 [64,65].
  • The PCMZ-3 zone is defined between 18.00 and 15.72 m. Based on the additional high proportion of thermophilous species, this zone also marks a warmer period, dominated by species preferring open vegetation (H. striata, P. triplicata, P. muscorum). C. tridens and V. costata are not significant at this level. Hygrophilous and cold-tolerant Succinella oblonga (17.04–16.80 m, in negligible proportions) appeared at one level, perhaps confirming a minor cooling period. The zone was characterised by a warm, dry period, based on the calculated temperature and humidity curves, apart from the S. oblonga level. The zone includes the upper part of the PS3 paleosol, the PL3 loess layer, and the lower half of the PS2 paleosol layer, so that correlations suggest an age of the zone between 280 and 220 kys, which corresponds to MIS sections 8–7e [64,65].
  • The PCMZ-4 zone represents the section between 15.72 and 12.96 m. The part of this zone in the paleosol PS2 is dominated by the shade-loving, wide-tolerant Limacidae species. Further along the zone, the proportion of Limacidae species decreases, with the wide-tolerant V. pulchella and P. muscorum and the thermophilous H. striata and P. triplicata appearing in smaller proportions. Entering the PL2 loess layer, the proportion of thermophilous and mesophilous species is approximately equal. In this layer (14.28–14.16 m), the cold-loving Columella columella, which prefers open vegetation, is present. Its presence and low proportion may indicate a minor cooling period. In the upper part of the zone, the high proportion of thermophilous and open vegetation-preferring species (especially H. striata) indicates a dry, open vegetation environment. The age of the zone can only be estimated by correlation, which ranges between 220 and 150 kys, covering the MIS 7 and part of the MIS 6 period [64,65].
  • The short section of the PCMZ-5 malaco-zone (12.96–9.48 m) on PL2 loess layer is characterised by warmth-loving dry, open vegetation preferring Granaria frumentum, H. striata, P. triplicata and cold-resistant S. oblonga (only between 12.96 and 12.84 m) preferring humid, open vegetation. The distribution of the faunal composition indicates periodic moister phases, which are also reflected in the humidity graph. The shade-loving species Clausilia dubia and Nesovitrea hammonis, which are wide-tolerant, and Discus ruderatus, a cold-resistant species that prefers coniferous forests, are present in the PS1 layer of the zone. Their distribution in the zone (as well as in PS1) reveals three forest expansion events between 12.72 and 12.48 m C. dubia, between 1092 and 1056 cm N. hammonis and D. ruderatus, where coniferous trees were probably present, and between 9.96 and 9.72 m again marked by C. dubia. At the latter two levels, species preferring open vegetation appear in addition to shade-loving species, suggesting a mosaic character of the landscape, i.e., the area may have developed into a steppe-like environment with trees. Temperatures estimated in the zone, due to the relatively high number of sterile samples, were quite variable, typically lower in the forest cover levels. The age of the zone can still be correlated with the age of the PS1 paleosol falling within MIS 5, i.e., between 127 and 73 kys [64,65].
  • The PCMZ-6 malaco-zone (9.48–6.00 m) already has absolute age data taken from the calculated age–depth model (from 8.84 m). The snail fauna of this zone is similar to PCMZ-2, but with a higher species richness and a higher diversity, especially of the cold-resistant and cryophilous species. Among the thermophilous species, the presence of open vegetation preferring species (H. striata, P. triplicata, C. tridens; G. frumentum has disappeared from the zone) dominates, but the proportion of mesophilous species is higher. In contrast to the predominance of P. muscorum, which prefers an open, dry environment, and V. costata, which prefers open vegetation, in the middle part of the zone (between 8.16 and 6.24 m in PL1L2 layer, ca. 45,714–38,579 cal BP), moisture-loving species such as Vallonia pulchella and V. enniensis, Euconulus fulvus and Limacidae species, as well as the cold-resistant S. oblonga (between 816 and 744 cm; 45,741–42,745 cal BP) occur, indicating a lower cooling period. The age of the zone is 49,172–33,235 cal BP, which falls within the MIS 3 interglacial period [64,65]. The moister period falls within the Greenland Stadial (GS) 9–12 and Greenland Interstadial (GI) 9–12 periods [66], i.e., covering the period of rapid alternation of stadials and interstadials.
  • The PCMZ-7 zone (6.00–4.68 m) ranges from 33,235 to 25,359 cal BP, covering the MIS 2 stage [64,65], including the GS 6-3 and GI 5.2-3 periods [66]. The total length of the stadials is much longer than the interstadials, indicating a colder period on average. The evolution of the malacofauna is also consistent with this. In addition to a significant decrease in the proportion of thermophilous species, there is an increase in the proportion of cold-loving species, in this case, Columella columella, which prefers cold, moist grassland. However, the proportion of C. columella is rather low (1–3%), but several cold-tolerant species also appear in the zone: the largest proportion being the shade-loving Trochulus hispidus, which prefers a more humid environment, and the moisture-loving S. oblonga. There is also a relatively low dominance of Pupilla sterrii, which prefers open, dry vegetation, and Discus ruderatus, which prefers coniferous forests. The absolute predominance of the mesophilous species is represented by P. muscorum and V. costata, but several shade-loving (Clausilia dubia, C. pumila, Nesovitrea hammonis, Euconulus fulvus, Limacidae) and hygrophilous (Vallonia enniensis, V. pulchella) species also appear. According to the composition of the snail fauna, the study area alternated between cool to moderately dry steppe and more humid forest areas, providing a habitat for a wide range of snail species. The average abundance per sample is the highest in the whole section, and diversity and species richness are also at their maximum here. Lower average temperatures and higher humidity resulted in a much more diverse malacofauna.
  • The PCMZ-8 zone was delineated between 4.68 and 2.40 m in the section, between 25,359 and 23,796 cal BP, falling entirely within the GS 3 stadial phase of the MIS 2 period [64,65,66]. The malacofauna of the Pécel loess–paleosol section shows a warmer and drier period during the stadial phase compared to the previous zone. The proportion of warmth-loving species was increased, with G. frumentum, C. tridens and Cochlicopa lubricella present in addition to the prominent proportions of H. striata and P. triplicata. All the thermophilous species present here prefer open vegetation, so the principal environment of the zone may have been dry grassland. Among the wide-tolerant species, P. muscorum prefers this environment, while the shade-loving species Euconulus fulvus, Punctum pygmaeum, and Clausilia dubia are also present in the lower part of the zone (468–396 cm; 25,359–24,965 cal BP). In addition to the cryophilous C. columella, Vertigo pygmaea, which also prefers open vegetation, was more abundant in the zone. The proportion of arboreal vegetation may have been higher in the first phase of the zone, but it gradually declined over time.
  • In PCMZ-9 (2.40–1.20 m; 23,796–22,235 cal BP), cold-resistant and cryophilous species disappear from the zone, indicating a rather warm and dry period, except for the earliest phase. The zone covers the GI 2.2 and 2.1 interstadials [66], but it cannot trigger such warming. Among the warmth-loving species, H. striata is extremely dominant, with P. triplicata also found subordinate. Among the wide-tolerant species, P. triplicata and V. costata have the highest proportion, and Limacidae, C. dubia and E. fulvus shells were also sporadically found. These species are evidence of the occasional occurrence of small groups of trees in the dry steppe area during this period.
  • PCMZ-10 (1.20–0 m) comprises the youngest malacofauna, from 22,235 cal BP, the GS 2.1 stadial period [66]. The latest calculated age date is 18,090 cal BP (44 cm). The snail fauna in this zone was also dominated by species preferring warm, dry, open vegetation (P. triplicata, G. frumentum, H. striata) and by the wide-tolerant P. muscorum and V. costata with similar ecological requirements. In the youngest part of the zone, the new species Vitrea crystallina, which prefers humid forests with wide tolerance, appears together with the cold-resistant Pupilla sterrii, Succinella oblonga and Trochulus hispidus. The co-occurrence of these species indicates a decrease in temperature and an increase in humidity, and thus the advance of arboreal vegetation.

5. Conclusions

Comparison of the magnetic susceptibility curves showed that the L1 layer could be identified with the Chinese Loess Plateau’s L1 layer, but the available 20 radiocarbon age data confirmed this and the correlation with the MIS 2–4 stages. The accumulation of the L1 loess body was presumably interrupted by an interstadial stage (L1S1), which also affected the accumulation rate of the lower part of the L1L1 section. The increased AR and MAR values in the L1L1 layer compared to the L1L2 section certainly indicate the increase in transport energy; moreover, they make possible the determination of the prevailing wind direction. In addition to the recent research approaches carried out in this paper, OSL studies could clarify the actual age of the lowermost part of the profile; however, the MS curve compared to the Chinese Loess Plateau’s curve allows the expression of relative ages within the entire loess–paleosol section. After interpreting the results of the malacological examination, some important conclusions can be drawn about the paleoenvironmental and paleoclimatic conditions of the Pécel loess–paleosol section. The temperature reconstructed on the basis of the malacological data was basically warm, even during the stadial periods, which can be explained by the southern exposure of the section. The predominance of species preferring open vegetation suggests that the vegetation cover was dominated by open steppe vegetation, with only a few levels showing significant forest expansions (mainly in the PS2, PS1 and PL1S1 paleosol layers). The appearance of shade-loving species typically resulted in an increase in humidity and a decrease in temperature (Figure 5), which had a positive effect on the diversity of the malacofauna. The constant presence of the thermophilous Helicopsis striata (Figure 6) in almost all the examined samples suggests a refugium of this species in the vicinity of Pécel, similarly to the refugium of Pupilla triplicata found in Villánykövesd (SW-Hungary) [61].

Author Contributions

Conceptualization, L.M., D.M. and P.S.; methodology, L.M. and D.M.; software, L.M. and D.M.; validation, L.M. and D.M.; formal analysis, P.C.; investigation, L.M., D.M., P.C. and B.N.; resources, L.M., D.M. and P.S.; data curation, L.M.; writing—original draft preparation, L.M.; writing—review and editing, L.M., D.M. and P.C.; visualization, L.M. and D.M.; supervision, P.S.; project administration, L.M. and D.M.; funding acquisition, L.M. and P.S. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Hungarian Government, Ministry of Human Capacities (20391-3/2018/FEKUSTRAT), the New National Excellence Programme grant ÚNKP-22-4-SZTE-503 (L. Makó) and the European Regional Development Fund, grant number: GINOP-2.3.2-15-2016-00009 ‘ICER’.

Data Availability Statement

The data presented in this study are available on request from the corresponding author. The data are not publicly available because they will also be used in ongoing research.

Acknowledgments

We are grateful to Gedeon Rajcsinec for the protection of the area, thus allowing the loess wall to survive and for giving us the opportunity for the investigations.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Location of the loess–paleosol profile of Pécel in the Carpathian Basin: 1. Buda Thermal Karst, 2. Börzsöny, 3. Cserhát [11].
Figure 1. Location of the loess–paleosol profile of Pécel in the Carpathian Basin: 1. Buda Thermal Karst, 2. Börzsöny, 3. Cserhát [11].
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Figure 2. Age–depth model derived from 20 radiocarbon data in Bacon software [29].
Figure 2. Age–depth model derived from 20 radiocarbon data in Bacon software [29].
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Figure 3. Accumulation Rate and MAR values derived from them. The highlighted layer is the paleosol, green line: minimum ages, blue line: average ages, red line: maximum ages.
Figure 3. Accumulation Rate and MAR values derived from them. The highlighted layer is the paleosol, green line: minimum ages, blue line: average ages, red line: maximum ages.
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Figure 4. Results of magnetic susceptibility measurements compared with the Chinese Loess Plateau’s data series [12].
Figure 4. Results of magnetic susceptibility measurements compared with the Chinese Loess Plateau’s data series [12].
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Figure 5. Analysis of the molluscs material of the loess–paleosol sequence of Pécel.
Figure 5. Analysis of the molluscs material of the loess–paleosol sequence of Pécel.
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Figure 6. Percentage distribution of snail species.
Figure 6. Percentage distribution of snail species.
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Table 1. Radiocarbon ages from the Pécel sequence [25,26].
Table 1. Radiocarbon ages from the Pécel sequence [25,26].
Lab CodeDepth (cm)Materialuncal. BPσcal. BPσProbability Distribution
DeA-3428812–24G. frumentum13812112971692%
DeA-34278108–120T. hispidus18,1055822,062159100%
DeA-34289132–144H. striata19,6986323,7817465%
DeA-34279264–276T. hispidus20,1356824,11923599%
DeA-34290276–288H. striata20,3616924,442240100%
DeA-34280300–312T. hispidus20,0046724,022184100%
DeA-34291336–348H. striata20,5566624,77326199%
DeA-34281360–372T. hispidus20,6347124,862244100%
DeA-34282408–420T. hispidus20,7757225,043220100%
DeA-34292432–444H. striata20,9027025,18318896%
DeA-34283468–480T. hispidus21,1137025,546124100%
DeA-34293480–492T. hispidus20,5416724,75125697%
DeA-34284516–528T. hispidus34,64020639,937593100%
DeA-34294540–552T. hispidus21,2667325,568245100%
DeA-34285552–564T. hispidus21,0087325,385232100%
DeA-34295576–588T. hispidus20,1416624,12623398%
DeA-34296624–636H. striata35,21421440,349519100%
DeA-34286648–660T. hispidus38,90432542,607277100%
DeA-34287744–756T. hispidus39,38533442,803343100%
DeA-34297756–768H. striata38,63929042,508248100%
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Makó, L.; Cseh, P.; Nagy, B.; Sümegi, P.; Molnár, D. Paleoecological Reconstruction Derived from an Age–Depth Model and Mollusc Data, Pécel, Hungary. Quaternary 2025, 8, 37. https://doi.org/10.3390/quat8030037

AMA Style

Makó L, Cseh P, Nagy B, Sümegi P, Molnár D. Paleoecological Reconstruction Derived from an Age–Depth Model and Mollusc Data, Pécel, Hungary. Quaternary. 2025; 8(3):37. https://doi.org/10.3390/quat8030037

Chicago/Turabian Style

Makó, László, Péter Cseh, Balázs Nagy, Pál Sümegi, and Dávid Molnár. 2025. "Paleoecological Reconstruction Derived from an Age–Depth Model and Mollusc Data, Pécel, Hungary" Quaternary 8, no. 3: 37. https://doi.org/10.3390/quat8030037

APA Style

Makó, L., Cseh, P., Nagy, B., Sümegi, P., & Molnár, D. (2025). Paleoecological Reconstruction Derived from an Age–Depth Model and Mollusc Data, Pécel, Hungary. Quaternary, 8(3), 37. https://doi.org/10.3390/quat8030037

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