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Article

First Records of Beetle Fauna (Insecta: Coleoptera) from Late Glacial Sediments of Lithuania: Novel Environmental Reconstructions

by
Nick Schafstall
1,*,
Miglė Stančikaitė
1,
Romas Ferenca
2 and
Vaida Šeirienė
1
1
State Scientific Research Institute Nature Research Centre, Akademijos St. 2, 08412 Vilnius, Lithuania
2
Tadas Ivanauskas Museum of Zoology, Laisvės av. 106, 44253 Kaunas, Lithuania
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(12), 820; https://doi.org/10.3390/d17120820
Submission received: 27 October 2025 / Revised: 20 November 2025 / Accepted: 24 November 2025 / Published: 27 November 2025
(This article belongs to the Section Biogeography and Macroecology)
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Abstract

This study presents the first subfossil beetle (Coleoptera) records from Lithuania, from Late Glacial organic deposits. Bulk sediment samples were collected from the Pamerkiai and Zervynos Outcrops in SE Lithuania, and from the Ventė Outcrop at the eastern coast of the Curonian Lagoon, W Lithuania. Radiocarbon dating determined that the studied sediments accumulated between ~15,000–11,300 cal BP. The beetle assemblages (29–177 individuals per sample) consist of many cold-adapted species that are common from Late Glacial deposits in the British Isles, Southern Sweden, and continental Europe. True arctic species are absent from the assemblages, and it is likely that the Lithuanian beetle fauna was most similar to nearby southern regions (e.g., Poland) during the Late Glacial. Besides a variety of aquatic species and typical wetland species, many beetle species living in open environments and on sandy soils were identified. In almost all the samples, taxa associated with pine trees, willows, and birches were found, confirming previous reconstructions of a sparsely forested landscape during the climatic periods GI-1e–GI-1a (Bølling-Allerød). The species assemblages from the youngest samples, associated with GS-1 (Younger Dryas), indicate the disappearance of large aquatic macrophytes and decreasing temperatures in Southern Lithuania, but a persistence of trees in the region.

1. Introduction

Paleoecology is a special field of ecology that studies the long-term features and development of species, landscapes, and ecosystems by deriving properties from biotic fossils from sediment records, or directly from the sediment. The reconstruction of the deep-time history of landscapes is of great importance so that we can better establish baselines to compare current ecosystems with. It also helps us to better understand the processes that have formed our landscapes and will continue to do so in the future. For a long time, plant pollen was the most frequently used proxy in palaeoecological research [1,2,3]. However, the study of macroscopic remains of plants and animals often reveals a greater number of details about the local environment and changes therein than is possible by palynology alone. In particular, the remains of beetles (Coleoptera) provide valuable insights into the paleo-environment. This is because beetle species do not adapt to their environment, but rapidly and continuously migrate in search of their preferred environmental, or climatic niche [4]. Many beetle species are stenothermic, adapting to a narrow range of temperatures, and can become established most rapidly in new regions in response to changing climates. Extensive research on glacial and interglacial deposits from the British Isles has revealed that there were specific groups of species that occurred during glacial, or interglacial periods [5]. The Pleistocene fossil record, for instance, suggests that certain species of beetles colonized newly deglaciated landscapes within a few years of the ice margins retreating [6]. Beetle faunas from the Late Glacial period are widespread across the whole of Northwestern Europe [7,8,9]. Despite more than 60 years of studies on Quaternary beetles, knowledge of subfossil beetle assemblages remains clustered in the British Isles [10]. Currently no research of this kind has been carried out in the Eastern Baltic region, even though the knowledge about fauna during the Late Glacial period remains limited.
The Last (Weichselian) Glaciation and the subsequent postglacial period experienced significant climate variations [11,12,13], during which the European landscape was shaped. This is especially true for Northern Europe, where the Scandinavian ice sheet persisted for the longest period. In the north, the retreating ice sheet left bare soils that soon were covered by aeolian dunes, as part of the European Sand Belt (ESB) [14,15]. Currently, the zone of the ESB consists of thick layers of sandy soils mostly covered by pine forests. The southern part of Lithuania became free of ice between ca. 18,000 and 15,000 cal BP according to 10Be dating of moraines [16], turning into undulated outwash plains formed by braided meltwater streams [17,18]. Investigations of Lithuanian Late Glacial aeolian deposits, which correspond to the northeastern part of the ESB, show that widespread wind-driven sedimentation began around 16,000–15,000 cal BP [19]. While most of the Late Glacial and Early Holocene sediments in Southern Lithuania are currently buried beneath several meters of aeolian sand, river valleys have exposed small outcrops of organic layers. These layers of organic matter were already recognized to be valuable for paleoecological research and have been studied to determine pollen, plant macrofossils, and diatoms [20,21,22,23]. Through our study on beetle remains from Lithuanian sediment outcrops, we present the first Quaternary entomological study from Lithuania and the whole Eastern Baltic region. Our results provide an insight into the diversity of the coleopteran fauna in this region during the Late Glacial period and the reconstruction of vegetation and landscape based on the ecological preferences of the identified species.

2. Materials and Methods

2.1. Site Descriptions and Sample Collection

Three outcrops were selected for this study, known to be of Late Glacial age and with recorded macrofossil content: the Pamerkiai and Zervynos Outcrops in Southeastern Lithuania, and the Ventė Outcrop in Western Lithuania (Figure 1).
The Pamerkiai Outcrop (54°18′45″ N, 24°43′52″ E) is located 500 m upstream from the village of Pamerkiai, in a bend of the river Merkys, in SE Lithuania. The locality was previously studied in the context of Late Glacial environmental development [21,22] and became also of interest in an archeological context when a Medieval timber construction was discovered in the river wall [23]. In the summer of 2023, a section around one meter thick was subsampled in 2 cm resolution to measure loss-on-ignition and grain sizes of the sediments. Here, the section of interest consisted of two distinctive gyttja (fine organic sediment) layers, interbedded with thick layers of sand (Figure 2). Material was collected from the top and bottom of both gyttja layers for radiocarbon dating. In the silty sand between the gyttja layers, a fine layer of ca. 5 mm of woody debris and charcoal was discovered (Figure 2 and Figure 3), of which ~100 mL was collected as a bulk sample. Finally, large bulk samples (ca. 10 L) were collected from both gyttja layers and the interlying sand layer of the subsampled section.
The Zervynos Outcrop is located in the valley of the Ūla River, ca. 25 km southeast of Pamerkiai, SE Lithuania. This thick sediment complex of fine silt and gyttja layers was previously studied for pollen, geochemistry, and plant macrofossils [20]. In the summer of 2024, the outcrop was exposed on the left bank of the Ūla River, about 250 m downstream from the Zervynos village railway station (54°06′29″ N, 24°29′12″ E). Here, the section consisted of three layers of 105 cm gyttja in total (Figure 2), interbedded with layers of coarse and medium-grained sand (grain size was visually estimated in the field). The top and bottom of each gyttja layer were sampled for radiocarbon dating. The bottom layer of gyttja was 55 cm thick and divided into three parts for bulk sample extraction (15; 15; 20 cm). Besides from the gyttja layers, one more bulk sample (ca. 10 L) was collected from a medium-grained sand (ca. 10 cm thick) situated between the top layer and the middle gyttja layer (Figure 2 and Figure 3).
The Ventė Outcrop is located in Ventės Ragas on the eastern shore of the Curonian Lagoon (55°21′02″ N, 21°12′06″ E) and is a coarse-grained sandy deposits that contains a thin layer of peat (Figure 2), previously dated as Late Glacial [26]. The site was visited in the spring of 2024, and the peat layer was exposed from the slumping sand. A desiccated layer of ca. 5 cm peat was discovered. A large bulk sample (ca. 10 L) was taken from the peat, and one smaller sample was taken for radiocarbon dating.

2.2. Radiocarbon Dating

Small samples (<100 mL) from the three outcrops were searched for plant seeds, preferably terrestrial plants but otherwise marsh plants such as Carex spp. In case not enough seeds were present in a sample, bulk sediment was collected. From the woody debris layer in the silt layer of the Pamerkiai Outcrop, both seeds and charcoal pieces were collected. All material was sent to the Vilnius Radiocarbon laboratory (Vilnius, Lithuania) for AMS survey. Radiocarbon dates were calibrated with the program OxCal using the IntCal20 atmospheric curve [27,28].

2.3. Insect Extraction

Of all three outcrops, bulk sediment samples were first soaked in hot water with 5% KOH to disaggregate the compacted gyttja or peat, after which the complete samples were wet sieved over two sieves with mesh sizes of 1 mm and 0.2 mm to divide between a larger and smaller fraction. The samples were first left to dry, then deposited in plastic buckets and soaked in paraffin oil, after which cold water was carefully poured into each bucket. Exoskeletons of insects are made of chitin that binds to paraffin oil [29]. The floating layer of paraffin oil, together with any organic remains that floated by themselves, was poured into a sieve, and detergent was used to extract the oil from the samples. Finally, the samples were rinsed with ethanol to drive out any soap remains. The two size fractions of floated material were collected in a medium of diluted ethanol, after which insect remains were picked out from each sample under a binocular microscope with 10–45× magnification.

2.4. Beetle Identifications and Environmental Classifications

Remains of Coleoptera (beetles) were glued on cards and identified by first comparing them to online picture collections (kerbtier.de; cassidae.uni.wroc.pl/Colpolon) and later to specimens of the beetle collection of the Tadas Ivanauskas Zoological Museum in Kaunas, Lithuania. Final identifications were made in the beetle collection of the Natural History Museum in London, United Kingdom. An effort was made to identify as many beetle remains as possible to the species level, or at least the genus level. Based on the number of heads, pronota, and elytra (sclerotized wing shields) found, the minimum number of individuals (MNI) of a genus or species was recorded. The ecological information from beetle taxa was collected from species descriptions from the website coleonet.de and summaries from the literature found in the database program BugsCEP [30]. A single ecological attribute was assigned to each beetle taxon and counted per MNI, to show a quantitative distribution of the general biotopes ‘Aquatic’, ‘Marsh’, ‘Open’, ‘Trees’, ‘Dung/Foul’, and ‘Indet’ (when a taxon could not be assigned a single ecological attribute, e.g., specimens were only identified to the genus level and different species have varying ecological attributes). The category Trees was only attributed to taxa when the taxon is with certainty dependent on trees, such as species feeding on tree leaves, or living under the bark of trees or in deadwood. In terms of an alternative analysis, Bugs Ecocodes [30]; based on Koch [31,32,33] were attributed to each identified taxon, regardless of the MNI of each taxon. Here, a taxon could have more than one Ecocode (e.g., Otiorhynchus nodosus has an Ecocode for Meadowland as well as Heathland and Moorland). The attribution of single ecotones to taxa gives a straightforward overview of the different landscapes present around the sample site and is useful to estimate, e.g., the dominance of the aquatic environment or the tree cover around the sample site. Based on a large suite of experimental work carried out in Britain [34], single ecotone reconstructions can be used to estimate forest cover by the percentage of facultative tree species and the density of herbivores by the percentage of dung-dwelling taxa (e.g., [35]). An alternative analysis of all potential Ecocodes, in turn, provides a more detailed description of the different (potential) habitats of the beetle taxa. (For example, Moorland; Meadow; Sandy/Disturbed for the generalized ecotone of open landscape. In this study, it also shows the difference between the share of taxa that are facultatively tied to trees and taxa that could be found in a forested landscape (e.g., shade-tolerant species.).

3. Results

3.1. Radiocarbon Dating

For the Pamerkiai Outcrop (PK), six AMS dates were received from the Vilnius Radiocarbon laboratory (Table 1, Figure 3). The woody debris layer from the sand between the two gyttja layers (Figure 3, Figure S1) contained plant seeds as well as pieces of charcoal, which were both dated. While the radiocarbon age of the plant seeds is similar to the top layer PK3, the charcoal pieces have a similar age to the bottom layer PK1. Calibrated ages show that the complex of the two gyttja layers and intersected sand layer was deposited between 13,746 and 12,500 cal BP. From the Zervynos Outcrop (ZV), eight AMS dates show that the gyttja layers were deposited between 15,540 and 11,336 cal BP. The top of layer ZV1 and the base of layer ZV2 show older dates than the base of the thick sediment complex of ZV1 and the top of ZV2. The sample from the Ventė Outcrop (VT) was dated back to 13,161–12,915 cal BP, which is comparable to previously published radiocarbon dates from this site [26].

3.2. Sequence Correlations

Analysis of the complete profiles of Pamerkiai, Zervynos, and Ventė in this study (Figure 3) shows the temporal correlation between the three sections. Based on the calibrated radiocarbon dates of the top and bottom of the gyttja layers, Pamerkiai (PK) and Zervynos (ZV) appear to alternate temporally. While ZV1 is constrained between ~15,000–14,215 cal BP, PK 1–3 are constrained within 13,320–12,500 cal BP (correlated with the sandy layer ZV3), and ZV4 between 12,609 and 11,336 cal BP. Ventė (13,161–12,915 cal BP) can be temporally correlated with PK 1–2 and ZV S3. It should be regarded, though, that the sandy layers PK2 and ZV3 could contain reworked, older material. This is certainly the case for ZV2, according to the 14C data. ZV2 appears to be an incursion of eroded sand within a mixed matrix of gyttja; this is quite plausible when considering that the bottom and top of ZV1 are almost the same age. Therefore, ZV2 was excluded from the analysis and ZV1 was treated as a single unit, with ages according to the bottom of ZV1 and top of ZV2.

3.3. Identifications and Ecological Classifications of the Beetle Remains

3.3.1. Ventė

The thin peat layer from the Ventė Outcrop produced a total of 144 individuals from 40 different taxa, from the families Carabidae, Dytiscidae, Gyrinidae, Hydrochidae, Helophoridae, Hydrophilidae, Staphylinidae, Scirtidae, Dryopidae, Byrrhidae, Cryptophagidae, Scarabaeidae, Chrysomelidae, and Curculionidae. Beetle remains were somewhat decomposed, but often unfractured preserved in the peat matrix. Aquatic and marsh-dwelling taxa comprise the majority of 80% of the assemblage. The complete list of identifications can be found in Table S1. Taxa in open landscapes form 12.5% of the assemblage (Figure 4), while tree-associated species and species living in dung or foul environments (e.g., rotting vegetation) form a very minor part.

3.3.2. Pamerkiai

Apart from a few well-preserved beetle remains in the layer of woody debris, layer PK2 contains only few insect macro remains and is unsuitable for quantitative analysis. Of the two gyttja layers, the bottom layer sample S1 contained more and somewhat better preserved remains than sample S3, but most insect remains from both layers were badly broken, and only pieces of individuals were found. Around 70% of the remains could be identified to the genus level and 40% to the species level; a total of 296 individuals from 99 different taxa. The complete list of identifications can be found in Table S1, and an overview of the shares of the different described ecotones is displayed in Figure 4 and Figure 5. PK1 and PK3 both consist of around 9% individuals of aquatic taxa, and 6–7% of taxa that are living in or on trees. Layer PK1 contains a majority (53%) of marsh-dwelling taxa and around 12% of taxa living in open landscapes. Close to 20% of the identified taxa could not be attributed to a single ecotone because they were only identified to the genus level; this group is defined as ‘Indet’. In layer PK3, the share of marsh-dwelling taxa has decreased to ca. 27% and open landscape taxa has increased to 40%. The share of dung/foul dwelling taxa is ca. 3% in the sample. The samples from Pamerkiai are more diverse than Ventė, with more different taxa in PK1 and PK3. From the potential ecotones for each taxon (Figure 5), ‘Trees’ is more frequent in the samples from Pamerkiai than from Ventė. In layer PK3, the ecotone ‘Sandy’ is more frequent than in PK1.

3.3.3. Zervynos

From the three remaining samples from the Zervynos Outcrop, a total of 228 individuals were identified from 92 different taxa. The complete list of identifications can be found in Table S1. Aquatic and marsh-dwelling taxa comprise 33–43%, tree-dwelling taxa 5–8%, dung/foul 3–6%, and open landscape taxa 26–40%. Compared to layer ZV1, the share of open landscape taxa increases 50% in ZV3. This share remains almost similar in ZV4, but the share of marsh-dwelling taxa decreases from 29% to 16%. The taxonomic diversity of the Zervynos samples (Figure 5) is similar to Ventė, but the number of taxa that are potentially associated with trees is higher. The number of taxa potentially associated with sandy/disturbed environment is particularly high layer ZV3.

4. Discussion

4.1. Cross-Site Correlation of Beetle Taxa

A detailed comparison of the species that occur at multiple sites, should contribute to our understanding of the commonalities and differences between Ventė, Pamerkiai, and Zervynos. Many of the species found in the Late Glacial deposits are currently rare or absent in Lithuania [36], but occur elsewhere in the Eastern Baltic region or in Scandinavia. The assemblages are not very similar to each other ([Table S2], [37]). Only a few beetle taxa identified to the species level occur at all three sites (Table 2): Olophrum consimile, O. fuscum (Staphylinidae), Arpedium brachypterum (Staphylinidae), Tachinus elongatus (Staphylinidae), Cytilus sericeus (Byrrhidae), and Otiorhynchus nodosus (Curculionidae); these species would likely have occurred in the wider region and are typically all indicators for an open, marshy landscape. Looking at individual pairings (VT-PK; VT-ZV; PK-ZV), there are more common beetle taxa between sites. Ventė has a few carabids, dytiscids, and staphylinid species in common with Pamerkiai, especially with sample PK1. Patrobus atrorufus and (especially) Pterostichus oblongopunctatus could indicate a nearby presence of trees, while P. nigrita/rhaeticus, Agabus sturmii, Gymnusa brevicollis, and Pselaphus heisei emphasize the presence of thick layers of litter and organic debris in a wet environment. Common species between Ventė and Zervynos are Arpedium quadrum (VT-ZV1–3), Tournotaris bimaculatus (VT-ZV1), and Dryops griseus (VT-ZV4). A. quadrum could represent a dryer, open component in the landscape, similar between Ventė and Zervynos but, perhaps, absent at Pamerkiai. D. griseus could be indicative for the presence of temporary fen pools at both VT and ZV4; while this landscape feature was likely present at the coast it is peculiar that it might have been present at Zervynos around 12,500–11,300 cal BP. T. bimaculatus (VT-ZV1) would be indicative for either a wetland or a riverbank with large macrophytes such as Phragmites or Typha, and the absence of this species at Pamerkiai and younger samples of Zervynos, provided it was a real absence and not a gap in the fossil records, might be another clue about the landscape and vegetation structure of during the time these gyttja layers were deposited.
There are several species discovered both at Pamerkiai and Zervynos. Pycnoglypta lurida (Staphylinidae) appears quite frequently in the samples aged ~13,000–11,300 cal BP (PK1, PK3, ZV3, ZV4), while Bledius spp. (Staphylinidae), Bromius obscurus, Phratora vitellinae (Chrysomelidae), Psammoporus sabuleti (Scarabaeidae), Protapion sp. (Brentidae), Ips sp., and Hylurgops palliatus (Curculionidae: Scolytinae) were present at both sites but less abundant. Besides P. lurida, all these species are common throughout Europe [31,32,33]. The species assemblage is representative of an open riverbed, with willows or poplars and conifer trees in the near distance. While a generalized reconstruction of the potential biotopes at Ventė (Figure 4) highlights the dominant components of aquatic and marsh-dwelling taxa, as well as an abundance of taxa that require open landscape, the detailed reconstructions for Pamerkiai and Zervynos (Figure 5) show a more nuanced result. Considering the proportion of each Ecocode, it appears that there were more taxa that potentially live among trees at Pamerkiai than at Zervynos. However, the proportion of taxa that only live on trees is almost similar between the two sites (Figure 4). The higher proportion of tree- or forest-associated taxa in the Pamerkiai assemblage results from a larger number of taxa, mainly Carabidae, that are not only moisture-loving but also shade-loving and, therefore often found in forested areas. However, a shaded biotope could also result from thick herb layers or uneven or rocky terrain. Since the distance between Pamerkiai and Zervynos is only 25 km, this highlights how the Late Glacial ice sheet foreland produced a heterogenous landscape. An important similarity between the two youngest samples of the two deposits is the occurrence of Agabus arcticus, which lives at higher latitudes in clear mountain lakes and Sphagnum pools [38].

4.2. Late Glacial Environmental Reconstructions Based on the Beetle Records

According to the AMS 14C dates, samples from the different outcrops are related to the period ~15,000–11,300 cal BP. The examined gyttja and sand layers can tentatively be correlated to the different climatic phases GI-1e –GS-1 (Bølling—Younger Dryas) [39] and where possible, we made a reconstruction of vegetation, landscape structure, and indicators of warmer or colder atmospheric temperatures and compare our results with existing records from Lithuania and the wider region.
GI-1e (14,692–14,075 cal BP)
Layer ZV1 from Zervynos, which was deposited in a dynamic environment (according to non-consecutive 14C dates from different parts of the layer which cover, however, only a short time interval) was deposited during this warmer climatic interval and comprises a large part of it.
ZV1 mostly contains taxa living in either wetlands (Stenus spp., Omalium caesum, Olophrum spp.) or open environments (Arpedium quadrum, Arpedium brachypterum, Bromius obscurus, Protapion sp.). Wetland vegetation was inhabited by Donacia spp., Plateumaris sericea, Bagous sp., and Tournotaris bimaculatus, and the vegetation was likely quite dense. The taxa occupying open areas indicate drier grassland or moorland near the wet depression, and A. brachypterum is regarded as a species of the arctic and alpine tundra [40]. There are, however, abundant indicators for the presence of trees near the site. The longhorn beetle Prionus coriarius is a reliable indicator for trees or mixed woodland as its larvae develop in a variety of deciduous trees, and more rarely, conifer trees [41]. Weevils (Curculionidae) associated with trees were found as well; Xylechinus pilosus colonizes mainly dead pines [42], Ips sp. colonizes dead or dying conifers, and Anthonomus phyllocola and Brachonyx pineti feed on pine needles. Given the low share of 8% of facultative tree taxa and a high share of taxa living in open landscapes, there was likely no dense forest present nearby but rather scattered trees. To our knowledge, this record is the first evidence of presence of pine trees growing in Southeastern Lithuania during GI-1e. Although there appears to be disagreement about the abundance of conifer trees in the Baltic region during GI-1e (14,692–14,075 cal BP), which could be due to regional differences [43], pine trees were already growing in Eastern Latvia around 14,300 cal BP [44].
GI-1d (14,075–13,954 cal BP)
This cold event, also known as the Older Dryas, might be captured in layer ZV3 (14,215–12,048 cal BP), but this sandy layer covers a period of almost 2000 years, which makes it impossible to distinguish this period in the beetle record.
Due to the period of 2000 years between the two gyttja layers ZV1 and ZV4, it is impossible to contribute sandy layer ZV S3 to a single climatic episode. Compared to the other samples from Pamerkiai and Zervynos, it has a high proportion of taxa living in open landscapes, such as Bledius spp., Pycnoglypta lurida, Otiorhynchus nodosus, and Psammoporus sabuleti. The weevil Sitona macularius can be found in sunlit dry places [33]. Chrysomelids and curculionids that feed on large marsh plants are absent, and the identified chrysomelid Donacia obscura feeds on sedges. On the river shores, open landscape taxa form the majority of this sample (40%) and includes many Staphylinidae species, such as A. quadrum, A. brachypterum, Acidota quadrata, and Bledius spp. Geodromicus plagiatus is widely distributed in Europe, although concentrated in mountain areas and the boreal zone, where it is found under rocks and between moss. The carrion beetle Silpha tristis is found in rotting vegetation and cadavers in grasslands and sandy dunes. The sand-dweller P. sabuleti was also present, and a piece of elytron from the green tiger beetle Cicindela campestris was found; this species is often described to be restricted to sandy ground. There were only three species related to trees identified. Two species of Cryptocephalus were found: both C. bipunctatus and C. punctiger are polyphagous on trees and bushes but are mostly related to birch and willow. Finally, the conifer bark beetle Polygraphus sp. colonizes dying pine and spruce trees [42]. Overall, the species assemblage represents a tundra climate where trees were still present, but it was not possible to determine whether the sample represents climatic episode GI-1d, GI-1b, or a mixture of episodes.
GI-1c (13,954–13,311 cal BP)
This period, which represents a warmer interval, is covered by layer PK1 from Pamerkiai (13,746–13,120 cal BP), especially as the calibrated top age of this layer ranges between 13,320 and 13,120 cal BP.
The aquatic and marsh-dwelling beetle community, which forms the absolute majority in this sample, appears to be very diverse. Of the identified aquatic species, Agabus sturmii prefers standing water but can also be found in slow-flowing, vegetated parts of streams and rivers. Ochthebius minimus occupies silty and muddy sides of ponds or streams with well-developed vegetation [45], while Limnebius crinifer prefers flowing water. On the contrary, Hydrobius fuscipes occupy stagnant pools with detritus or vegetation. Cyphon spp. is the most abundant wetland taxon in this assemblage (35 MNI) and is often also found on swampy banks and in leaf litter. The different taxa that can typically be found in wetlands indicate both dense vegetation or flood debris, and a more open environment. Olophrum consimile occurs on the edges of lakes and streams with abundant vegetation, P. lurida is an Eastern European species that inhabits wet moss and litter under wood or bushes [31,46], and several taxa were identified that typically live in rotting plant material, such as Corticarina spp., Megasternum obscurum, and Tachinus elongatus. On the other hand, O. nodosus feeds on various herbs in various wet to moderately wet, open biotopes [47]. Coelositona cinerascens feeds on Lotus spp., which mostly grow in moist open areas. The small scarab beetle P. sabuleti lives on sandy shores of lakes and rivers. Perhaps, also this sample represents a dynamic riverine environment with dense bank vegetation alternating with bare sandy soils. In terms of climate, it is important to note that besides the alpine tundra species A. brachypterum, also several individuals of A. quadrata were identified; this species occurs only at high latitudes over the Palearctic (or isolated in mountain areas) [31]. Despite a possible tundra environment, trees were still spread across the landscape, or maybe more typically, growing in the floodplain of the river. The chrysomelids Plagiodera versicolora, Phratora vitellinae, and Crepidodera sp. all feed on leaves of willow [33]. Laemophloeus muticus lives under the bark of fallen tree trunks of mainly birch, but also alder, aspen, and oak. Species living on conifer trees were found as well; Hylurgops palliatus and Orthotomicus suturalis both colonize dying conifers, where O. suturalis prefers pine over other conifer species. Perhaps, by this time the landscape had become more colonized by trees, despite the previous cold episode and lower temperatures compared to GI-1e. A previous study on the Pamerkiai Outcrop, that included plant macrofossils [22], found evidence of spruce, pine, and birch, but not of willows. It should be considered if the willows present were rather shrub-like and not large trees. Nevertheless, the presence of willows during GI-1c gives new information for this region.
GI-1b (13,311–13,099 cal BP)
This short cold climatic interval is probably represented by the beetle assemblage from the thin peat layer from Ventė (13,161–12,915 cal BP). This site is located on the coast of the Curonian Lagoon which, at that time, was in close proximity to the Baltic Ice Lake [48].
The beetle community is primarily aquatic and living in wetlands and represents a wetland rather than a riverine environment. While it was found that there is some similarity between species of Ventė and Pamerkiai S1, different species are more dominant. The aquatic community consists of Dytiscidae, Gyrinidae, Hydraenidae, Hydrophilidae, and Dryopidae. From the dytiscids, Ilybius fenestratus is the most numerous species, while Hydroporus rufifrons, Hydroporus angustatus, Agabus sturmii, and Colymbetes sp. are present in lower numbers. All these species, as well as Gyrinus substriatus, Hydrochus brevis, and Enochrus spp., are indicative of stagnant water with abundant vegetation. Besides phytophagous species in wetlands, such as Cyphon spp., Donacia aquatica, and T. bimaculatus, many species were found that would have been hunting on the vegetated shores of the wetland, e.g., P. nigrita/rhaeticus, Agonum micans, A. quadrum, and Gymnusa brevicollis. From the taxa living in open landscapes, A. brachypterum was the most abundant; together with A. sturmii indicators of a colder climate. One unique find at this site is an individual of Agoliinus (Aphodius) lapponum; this species is foremost associated with dung from herbivores. This species is currently found at higher latitudes [49] and might be another indicator of colder climate. Although there are indicators for a colder climate, the difference in local environment complicates a comparison between Ventė and Pamerkiai S1 and it cannot be concluded that a colder climate than during GI-1c is represented here. Although much less definite than in Pamerkiai S1, there are also some indicators for the presence of trees found here: Patrobus atrorufus, often found in damp woods but also between shrubs, Pterostichus oblongopunctatus, a species that can occur both in deciduous and conifer forest, and the conifer bark beetle Orthotomicus sp., which colonizes dead or dying conifer trees.
GI-1a (13,099–12,896 cal BP)
This period, which was the end of the warmer climatic episode that started as GI-1c, is most likely represented by sandy layer PK2 from Pamerkiai (13,120–12,730 cal BP).
This layer contained very few beetle remains, but the share of wetland beetles is significantly lower than in layer PK1. The majority of taxa in this sample consists of species associated with an open moist environment, such as the carabid Elaphrus riparius, the pill beetle Cytilus sericeus, and the tundra species A. brachypterum. However, many taxa are more associated with a sandy, disturbed habitat: Amara eurynota, Bledius spp., P. sabuleti, and Otiorhynchus ovatus. Archarius salicivorus, Brachonyx pineti, and H. palliatus indicate the presence of willows and pine trees. An important finding in this layer was the charcoal layer, dated at 13,585–13,341 cal BP (11,601 ± 49 14C); this would be, to our knowledge, the oldest dated charcoal in Lithuania.
GS-1 (12,896–11,703 cal BP)
This cold period at the end of the Late Pleistocene, commonly known as the Younger Dryas in the Northern Hemisphere, is represented by layer PK3 from Pamerkiai (12,730–12,500 cal BP). But also layer ZV4 from Zervynos (lower boundary 12,609–12,048 cal BP; upper boundary 11,822–11,336) was deposited during this climatic episode.
At Pamerkiai, other species were found that appear to have been absent from the older layers. Agabus arcticus, Gyrinus marinus, Gyrinussubstriatus and Helophorus minutus are all aquatic species that prefer or tolerate stagnant water, even though a specimen of Elmis sp. was identified as well. Coelostoma orbiculare, Cercyon tristis, Paederus riparius, Philonthus nigrita, and Donacia cinerea do not require dense bank vegetation. Taxa living in open areas dominate this sample with 40% of the assemblage. Newly identified are Calathus ambiguus and Calathus fuscipes, which are associated with open, dry biotopes, Bledius spp., Hippodamia sp. (Coccinellidae), B. obscurus, and Gymnetron sp.; also other species such as O. nodosus appear again in this sample. For the tree-associated species (ca. 7%), several taxa appear newly in this sample. Xylodromus testaceus is a rare species that is found under bark and in decaying wood of deciduous trees. Chrysomela sp. is associated with willow or poplar, and Anoplus plantaris larvae mine the leaves of birches. Also, taxa associated with conifer trees were found, such as Hylurgops glabratus and Ips sp. One specimen of Aphodius sp. was found, but it could not be determined if this was a facultative dung-dwelling species or rather a species that also lives in rotting vegetation.
At Zervynos, the number of marsh-dwelling species decreases significantly while aquatic taxa and tree-dwelling taxa increase (however, tree taxa do not increase compared to ZV S1). According to tree-dependent taxa (Phratora vitellinae, H. palliatus, Phloeostiba lapponica), willows and conifers were present around this period. It is important to note that the evidence for the presence of tree species throughout this period confirms tentative conclusions about trees persisting in the Eastern Baltic region in other studies [21,50]. Also in this sample, the aquatic beetle A. arcticus appears. The carabid Blethisa multipunctata was found exclusively in this sample; this species is common throughout Europe, where it displays an amphibious lifestyle at moderately vegetated ponds and shores [31,51]. In Lithuania, this species is considered as a glacial relict [36]. Among the species living in open environments (38%) the same species of Staphylinidae were found in previous samples of Zervynos, except for the tundra species A. quadrata. The presence of Bledius spp. in particular indicates an open, wet environment. The dermestid beetle Dermestes murinus makes a single appearance in this sample; this species can be found on cadavers in any outdoor environment [32]. Besides this carrion beetle, one specimen of the dung-feeding taxon Onthophagus sp. was identified. All species in this genus are associated with ungulate dung. Although this provides clear evidence for the presence of grazing animals, a herd of animals would likely have resulted in a much higher percentage of dung beetles in the species assemblage. In any case, osteological data proves the presence of mammals, i.e., Rangifer tarandus (reindeer) and Mammuthus primigenius (mammoth) in the Baltic region during the final stages of the Late Pleistocene [52,53].

4.3. Common Beetle Taxa in Europe During the Late Glacial

Most fossil records of Coleoptera are from Britain and records from other parts in Europe are quite rare. However, it is still worthwhile to explore the species found in this study in the existing fossil record. For instance, P. lurida, which is currently restricted in its distribution to Scandinavia and Finland (Figure 6), has been found in various Pleistocene deposits in Britain [54], Sweden [55], France (La Grande Pile in samples representing MIS3; [56]), Germany [57], and Poland [58] and appears to be a common species throughout Europe during the colder and cooler epochs, at least from MIS13 [59] until the early Holocene, even in montane areas of Italy [60]. Interestingly, the species was regularly found together with Olophrum fuscum, Arpedium brachypterum, Aphodius lapponum, Otiorhynchus nodosus, and Tournotaris bimaculatus and at some sites also with Patrobus assimilis and Acidota quadrata. It appears that the more common species from the Southeastern Lithuanian deposits were generally common throughout Europe and the UK, occupying much wider areas during colder or cooler climatic episodes. Some species that are rare in the Lithuanian assemblage, for instance Bembidion assimile, are also rarer in the fossil record, and this carabid appears to have been a more western-oriented species as well during the Pleistocene and Late Glacial, with records only from Britain, the Netherlands, and France. The limited dispersal of this species might be related to the fact that individuals are frequently brachypterous [61].
Lemdahl [66] published an overview of the Late Weichselian and Early Holocene beetle faunas in Southern Sweden, in which he combined results from 30 sites to track occurrences of arctic species and more thermophilus species, or tree-dwelling species. This was further summarized in [67]. Southern Sweden became deglaciated around 15,000 cal BP, which was later than the deglaciation in Southern Lithuania. From deposits of this age, A. brachypterum and Agabus acrticus were found, together with, e.g., Apion spp. that indicate an open landscape in an early stage of succession. During the climatic episode GI-1e-1a (Bølling-Allerød), many species similar to the Lithuanian assemblage occurred. Pycnoglypta lurida, Olophrum consimile, A. brachypterum, Acidota crenata, A. quadrata, and Lesteva longoelytra, as well as Cyphon spp., were assumed to dwell in the leaf litter of willow and birch shrubs; it is an important observation that these trees might have only been present in shrub form. Donacia spp. were living on Carex and Scirpus, and O. nodosus and Aphodius spp., typical indicators of open landscapes, were common as well. But also, the more thermophilus carabid Pterostichus strenuus was frequently found in organic sediment deposits of this age. A reoccurrence of arctic species in S Sweden during GS-1 (Younger Dryas, 12,900–11,700 cal BP) was interpreted as an advance of the ice sheet to the south. More thermophilus species appear only in S Swedish deposits from the beginning of the Holocene. While the beetle assemblages from the Lithuanian deposits are quite similar with respect to the Southern Swedish fauna during GI-1e-1a, they contain many more thermophilus or late-successional species, such as Pterostichus oblongopunctatus and Calathus spp. Laemophloeus muticus, identified from sample PK1, which are currently associated with old-growth forest [36] and their presence suggests the presence of tall trees within several hundred meters from the stream channels. Another strategy to test the successional stage of a landscape, often used in studies on areas that were newly colonized after a retreating glacier (e.g., [68]), is the presence of brachypterous species, mostly from the family Carabidae, which would be the last to colonize an area due to their limited dispersal capabilities. The only macropterous species found in this study, Carabus nemoralis, was found in sample ZV1. In this context, the landscape might have been already well-developed during GI-1e. During the last cold interval before the onset of the Holocene, GS-1, the Lithuanian assemblage indicates that despite a climatic deterioration, trees still persisted in the landscape; this is an important difference compared to Southern Sweden.
Among the limited number of Easter European sites with common ‘Late Glacial’ or cold-adapted species, the Western Polish site Zabinko [58] is of special interest. A. arcticus, another common species in Late Glacial deposits, including the Southeastern Lithuanian sites in this study, is here found together with P. lurida, A. brachypterum, and O. nodosus. One of the conclusions of Lemdahl [58] was that the insect assemblage reflects an earlier deglaciation than in Fennoscandia, and that the landscape already started to develop during the Late Glacial. In this respect, the timing of ice sheet retreat, as well as the development of the landscape and climatic ameliorations in (Southeastern) Lithuania, would likely have been more comparable to Northern Poland than to Southern Sweden.

5. Conclusions

The records of Coleoptera from this study provide novel information that improves the current knowledge about the Late Glacial environment of Lithuania and the whole Eastern Baltic region. Indicator beetle taxa show that conifer trees were present in Southeastern Lithuania during GI-1e. Later, during GI-1c-1a, scattered tree communities on the glacier outwash plain consisted of pines, willows, and birches (possibly in shrub form). A beetle record from the peat sample from Ventės Ragas, which could be correlated to GI-1b, suggests a decline in trees during this climatic event. The onset of climatic episode GS-1 (Younger Dryas) is characterized by a decrease in wetland species and an increase in species living in open landscapes. A charcoal layer from the Pamerkiai Outcrop, dated 12,886–12,726 cal BP, proves aridity of the climatic regime. Agabus arcticus, found in both Pamerkiai and Zervynos, confirms that the climate deteriorated and that large aquatic macrophytes locally disappeared during GS-1. Arctic species were absent from all the samples, but typical tundra species were found in the samples, including the ones that are likely to represent warmer climatic episodes. The beetle assemblages from Ventė, Pamerkiai, and Zervynos do not only provide additional Supporting Information about the paleoclimate and landscape of Lithuania, but also directly provide insight in how the entomofauna responded to the climatic fluctuations and landscape developments during the Late Glacial. The abundant species from the Lithuanian sites are also common species in Late Glacial deposits from the European mainland. Besides that, an absence of arctic species suggests that climatic and ecological developments after the last glaciation follow patterns from, e.g., Poland. The records in this study, which contain various cold-adapted tundra species that currently live in montane regions or Scandinavia, are a valuable addition to Western European records to reconstruct the past, present, and future distribution of insect communities.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/d17120820/s1. Table S1: Complete list of species; Table S2: Bray–Curtis dissimilarity values; Figure S1: PK grain size analysis.

Author Contributions

Conceptualization, N.S., M.S., and V.Š.; Methodology, N.S.; Software, N.S.; Validation, N.S.; Formal Analysis, N.S.; Investigation, N.S.; Resources, R.F.; Data Curation, N.S.; Writing—Original Draft Preparation, N.S.; Writing—Review and Editing, N.S., M.S., R.F., and V.Š.; Visualization, N.S.; Supervision, V.Š.; Project Administration, N.S.; Funding Acquisition, N.S., M.S., and V.Š. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Lithuanian Research Council under grant number (P-PD-23-192).

Data Availability Statement

All data is available in the figures, tables and Supplementary Material of this article. Beetle Ecocodes can be accessed in the MS Access program BugsCEP [30] or online though SEAD (sead.se).

Acknowledgments

We would like to thank Vytautas Tamutis, Max Barclay, Dmitry Telmov, Alexey Shavrin, and Genrik Davidian for their help with identifying some of the subfossil beetle remains to the genus or species level. We also want to thank Oleksiy Davydov, Albertas Bitinas, Andrejus Petrovas, and Michal Šujan for their help during the fieldwork. Michal Šujan discovered the wood/charcoal layer during the sampling of the Pamerkiai Outcrop and shared a template from his own work which was used to create Figure 2.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Paleogeographical situation during the final stages of the Late Weichselian Glaciation in the territory of Lithuania (compiled by R. Guobytė [24]; Lietuvos nacionalinio atlaso žemėlapių rinkinys (I tomas) © Nacionalinė žemės tarnyba prie Žemės ūkio ministerijos, 2014) with identification of the ice-marginal formations [25]).
Figure 1. Paleogeographical situation during the final stages of the Late Weichselian Glaciation in the territory of Lithuania (compiled by R. Guobytė [24]; Lietuvos nacionalinio atlaso žemėlapių rinkinys (I tomas) © Nacionalinė žemės tarnyba prie Žemės ūkio ministerijos, 2014) with identification of the ice-marginal formations [25]).
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Figure 2. The three sampled outcrops, Pamerkiai, Zervynos, and Ventė, with the sampled layers marked.
Figure 2. The three sampled outcrops, Pamerkiai, Zervynos, and Ventė, with the sampled layers marked.
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Figure 3. Stratigraphic profile of investigated outcrops with sand and gyttja layers. Radiocarbon dates (cal BP) are shown as stars. Layer PK2 in the Pamerkiai Outcrop contained chaercoal fragments, while the sand above layer PK3 contained wave ripples. The lowest gyttja layer in the Zervynos Outcrop contained shells from mollusks and bivalves.
Figure 3. Stratigraphic profile of investigated outcrops with sand and gyttja layers. Radiocarbon dates (cal BP) are shown as stars. Layer PK2 in the Pamerkiai Outcrop contained chaercoal fragments, while the sand above layer PK3 contained wave ripples. The lowest gyttja layer in the Zervynos Outcrop contained shells from mollusks and bivalves.
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Figure 4. Share of generalized ecotones per sample, derived from the beetle assemblages. Only one ecotone was attributed to each taxon. Numbers between brackets show the number of individuals per sample.
Figure 4. Share of generalized ecotones per sample, derived from the beetle assemblages. Only one ecotone was attributed to each taxon. Numbers between brackets show the number of individuals per sample.
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Figure 5. Potential ecotones per sample, based on all related ecotones to the identified beetle taxa. Taxa can have more than one potential ecotone. Numbers between brackets show the number of taxa per sample.
Figure 5. Potential ecotones per sample, based on all related ecotones to the identified beetle taxa. Taxa can have more than one potential ecotone. Numbers between brackets show the number of taxa per sample.
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Figure 6. Discovered beetle remains from the Late Glacial Lithuanian outcrops that are currently absent in Lithuania but common in Scandinavia. For Pycnoglypta lurida (A), Arpedium brachypterum (B), Otiorhynchus nodosus (C), and Agabus arcticus (D), the subfossil remains identified in this study, a modern specimen, and the modern distribution according to GBIF [62,63,64,65] are shown. Red crosses indicate the locations from which the subfossil remains were identified.
Figure 6. Discovered beetle remains from the Late Glacial Lithuanian outcrops that are currently absent in Lithuania but common in Scandinavia. For Pycnoglypta lurida (A), Arpedium brachypterum (B), Otiorhynchus nodosus (C), and Agabus arcticus (D), the subfossil remains identified in this study, a modern specimen, and the modern distribution according to GBIF [62,63,64,65] are shown. Red crosses indicate the locations from which the subfossil remains were identified.
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Table 1. Results of AMS measurements from the Pamerkiai, Zervynos, and Ventė outcrops.
Table 1. Results of AMS measurements from the Pamerkiai, Zervynos, and Ventė outcrops.
Pamerkiai Outcrop
DepthSample nameMaterialLab codeAge + SD (yr BP)Cal. age (cal BP)CI
40 cmPK3 topPlant seedsFTMC-BC34-710,647 ± 4912,740–12,50095.4%
64 cmPK3basePlant seedsFTMC-LV69-1110,913 ± 5512,970–12,73995.4%
95 cmPK2Plant seedsFTMC-LV69-1010,849 ± 5212,886–12,72695.4%
95 cmPK2CharcoalFTMC-LV69-911,601 ± 4913,585–13,34195.4%
102 cmPK1 topPlant seedsFTMC-LV69-811,712 ± 5113,746–13,46295.4%
122 cmPK1 basePlant seedsFTMC-BC34-611,351 ± 4813,320–13,12095.4%
Zervynos Outcrop
DepthSample nameMaterialLab codeAge + SD (yr BP)Cal. age (cal BP)CI
0 cmZV4 topPlant seedsFTMC-LV69-2110,064 ± 5511,822–11,33695.4%
25 cmZV4 basePlant seedsFTMC-LV69-1810,426 ± 5112,609–12,04895.4%
35 cmZV2 topPlant seedsFTMC-LV69-1712,413 ± 6014,924–14,21595.4%
45 cmZV2 basePlant seedsFTMC-BC34-1013,865 ± 4717,020–16,64095.4%
50 cmZV1 topPlant seedsFTMC-LV69-1513,418 ± 4916,347–15,96295.4%
95 cmZV1 baseBulk sedimentFTMC-BC34-912,779 ± 4515,540–15,08095.4%
105 cmZV1 basePlant seedsFTMC-BC34-812,202 ± 5414,330–13,87095.4%
105 cmZV1 basePlant seedsFTMC-LV69-1411,751 ± 5113,752–13,50095.4%
Ventė Outcrop
DepthSample nameMaterialLab codeAge + SD (yr BP)Cal. age (cal BP)CI
270 cmVTPlant seedsFTMC-LV69-1311,135 ± 4913,161–12,91595.4%
Table 2. Common taxa from the three studied outcrops Ventė (VT), Pamerkiai (PK), and Zervynos (ZV). The minimum number of individuals (MNI) is shown, as well as a simplified ecotone for each of the taxa. Some taxa are only identified to the genus level.
Table 2. Common taxa from the three studied outcrops Ventė (VT), Pamerkiai (PK), and Zervynos (ZV). The minimum number of individuals (MNI) is shown, as well as a simplified ecotone for each of the taxa. Some taxa are only identified to the genus level.
Most common taxa (MNI)EcotoneVTPKZVTotal
Dytiscidae
Ilybius fenestratus (F.)Aquatic100010
Staphylinidae
Pycnoglypta lurida (Gyll.)Wetland017522
Olophrum consimile (Gyll.)Wetland22131549
Olophrum fuscum (Grav.)Wetland5319
Arpedium quadrum (Grav.)Open301215
Eucnecosum brachypterum (Grav.)Open851629
Stenus spp.Wetland26141252
Scirtidae
Cyphon spp.Wetland1638357
Byrrhidae
Cytilus sericeus (Forst.)Open1449
Latridiidae
Corticarina sp.Foul010010
Curculionidae
Otiorhynchus nodosus (Müll.)Open13610
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Schafstall, N.; Stančikaitė, M.; Ferenca, R.; Šeirienė, V. First Records of Beetle Fauna (Insecta: Coleoptera) from Late Glacial Sediments of Lithuania: Novel Environmental Reconstructions. Diversity 2025, 17, 820. https://doi.org/10.3390/d17120820

AMA Style

Schafstall N, Stančikaitė M, Ferenca R, Šeirienė V. First Records of Beetle Fauna (Insecta: Coleoptera) from Late Glacial Sediments of Lithuania: Novel Environmental Reconstructions. Diversity. 2025; 17(12):820. https://doi.org/10.3390/d17120820

Chicago/Turabian Style

Schafstall, Nick, Miglė Stančikaitė, Romas Ferenca, and Vaida Šeirienė. 2025. "First Records of Beetle Fauna (Insecta: Coleoptera) from Late Glacial Sediments of Lithuania: Novel Environmental Reconstructions" Diversity 17, no. 12: 820. https://doi.org/10.3390/d17120820

APA Style

Schafstall, N., Stančikaitė, M., Ferenca, R., & Šeirienė, V. (2025). First Records of Beetle Fauna (Insecta: Coleoptera) from Late Glacial Sediments of Lithuania: Novel Environmental Reconstructions. Diversity, 17(12), 820. https://doi.org/10.3390/d17120820

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