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Article

Wetlands in the Pleistocene Steppe-Tundra Landscapes of Beringia, Their Insects, and the Role of Aeolian Sedimentation

Laboratory of Arthropods, Borissiak Paleontological Institute, RAS, Profsoyuznaya 123, Moscow 117868, Russia
Water 2023, 15(3), 494; https://doi.org/10.3390/w15030494
Submission received: 3 December 2022 / Revised: 19 January 2023 / Accepted: 21 January 2023 / Published: 26 January 2023
(This article belongs to the Section Water and Climate Change)

Abstract

:
Analysis of the database of Beringian subfossil insect assemblages showed a relatively low role of aquatic, riparian, and wetland species of insects with hard exoskeleton in the Pleistocene communities and an increase in their proportions and taxonomic diversity in the Holocene. Aquatic insects were represented in all types of geological deposits and in some paleosols, but their proportions varied in different depositional environments. Poor representation of aquatic insects and a lack of freshwater invertebrates in the Late Pleistocene ice-rich deposits of Beringia called Siberian Yedoma or Yukon Muck attest to the predominantly aeolian origin of this phenomenon.

Graphical Abstract

1. Introduction

Aquatic insects played an immense role in continental ecosystems of the past. Most of the recorded fossil insects come from sediments of water origin [1]. Aquatic insects dominate many fossil assemblages; in some facies, they are extremely abundant and widespread. Among insects, only aquatic species are suitable for stratigraphic purposes: for example, the mayfly Ephemeropsis trisetalis (Eichwald, 1864) and the beetle larvae Coptoclava longipoda (Ping, 1928) are used for the stratigraphic correlation of the Early Cretaceous of Transbaikalia, Mongolia and North-Eastern China [1].
Quaternary insects that are relatively young and not fully fossilized could be found in various depositional environments from alluvial and lacustrine to loesses and paleosols. The proportion of aquatic species in Quaternary insect assemblages varies widely, from high to low to the complete absence. Almost all Quaternary insects are modern species [2]; they are not used in the traditional chrono-stratigraphy, which is based on evolutionary changes, but can contribute to the climate-based stratigraphic correlation. Aquatic insects provide important information on climate-related large-scale range shifts, local extinctions, and expansions [3]. For example, the fossil Helophorus orientalis Motschulsky, 1860 was found in the Pleistocene of Europe, far from its modern range that includes North America and Asia.
A unique example of a low evolutionary rate is the aquatic beetle Helophorus sibiricus Motschulsky, 1860. The well-preserved fossil from the Early Miocene of Siberia is morphologically identical to the modern H. sibiricus [4]; this is the oldest recorded evidence of a “living fossil” among hydrophiloid beetles and probably the only one properly documented case among all beetles.
Beetles can help in solving some specific geological tasks. The presence of aquatic insects in presumably windborne loess-like deposits, the low proportions of forest insects in “forest beds” and many other interesting discrepancies affect our assumptions about the depositional environments. That is of special importance for the long and emotional discussion of the origin of the ice-rich silty sediment called “Yedoma” (Figure 1a,b) in North-Eastern Siberia [5] and the Yukon Muck in North America.
This paper is focused on aquatic, riparian, and wetland insects with a solid chitin exoskeleton from the spacious ice-free land of Beringia (Figure 2). This land is stretched across the north-western part of North America and in the north-eastern part of Asia, including the Bering Strait (the Bering Land Bridge of the Pleistocene).
We aimed to describe subfossil insect assemblages specifically for various depositional environments in the region and further apply the association between deposition types and the insect assemblages to answer the question of the origins of the specific Late Pleistocene ice-rich sediments, Yedoma.

2. Materials and Methods

According to the classical taphonomy [6,7], almost all body fossils are preserved in aquatic depositional environments. This assumption is valid for true fossils [1] but relatively young late Neogene–Quaternary subfossil insects potentially can be found in deposits of various origins, including loesses and paleosols.
Insect remains of the Late Neogene and Quaternary are represented by the chitin devoid of soft tissues. The fossilization process of Beringian insects is not complete yet, because of the comparatively short period of time since their burial and due to the excellent preservation properties of the permafrost. These insect assemblages can be termed as taphocoenosis, an assemblage of dead organisms buried in the sediment without mineralization and other taphonomic changes.
In the Late Neogene and Quaternary deposits, we can find separated fragments of beetles, ants, true bugs (Hemiptera), and other hard chitin insects. In many cases the number of preserved fragments differs from their number in living insects: for example, we have recorded one head, two pronotums, ten left elytra and 20 right elytra in a single sample. This means that insect bodies were disassembled prior to the burial, and the fragments were transported and sorted by their size, shape, etc., during redeposition. However, in some cases, the number of fragments may coincide with that in the living organisms, and therefore, the Fragment Correlation (FC) can be of use as an indicator of the transportation mode and sedimentation type.
The degree of preservation is another variable. Permafrost provides excellent preservation of organic remains, but some of them may be broken and worn. Such damage may point to the long-distance transport of macrofossils prior to their redeposition. For instance, round stains and holes on chitin fragments are indicative of the long exposure of the fragments at the day surface prior to their burial (S. Kiselev, pers. comm.). Therefore, the reconstruction of depositional environments can be carried out by combining information on proportions of various ecological groups, number of fragments preserved, and even wear marks on insect macrofossils.

2.1. Lacustrine Deposits

Lakes were widespread in Beringia through the entire Quaternary period, including the recent time (Figure 3a).
Thermokarst lakes typically formed during interglacial intervals [8,9]. The thermokarst lakes start growing on ice-rich sediments as a result of climate warming. They accumulate organic matter from the eroding ground around the lake and from the thawing deposit beneath the lake. The thickness of the resulting lake deposits may reach several meters. Thawed ice wedges form small local depressions, ice wedge casts (Figure 1e) that can accumulate sediments of seasonal/temporary ponds or lacustrine sediments.
During cold climatic phases, some lakes were permanently covered by ice. These lakes are expected to be lifeless, with deposits lacking any organic inclusions. Subfossil insects and freshwater invertebrates are excellent indicators that allow one to identify such lakes. The absence of insect remains indicates that the lakes were lifeless indeed, but we know examples of some seemingly empty inorganic sediments that contained plenty of small chitin fragments. It means that water in the lake was open at least during the short summertime.
In cold climates, short-lived lakes often form in interdunal depressions. In a geological section, such sediment looks like a thin dark-colored layer bedded between sand deposits (Figure 1d). Wind may also create shallow depressions in the steppe that are filled by water during the spring season or even longer (Figure 4a). The cold climate may prevent complete desiccation of such a water body, but its volume would undergo dramatic seasonal changes. As we will see later, similar steppe lakes or “sloughs“ played an important role in the ecosystems of Beringia.
Lacustrine deposits are characterized by a high proportion of water insects, and the presence of shells of freshwater Gastropoda and Bivalvia (Figure 3b), statoblasts of freshwater Bryozoa (Phylactolaemata), Daphnia winter eggs (ephippia), Ostracod shells, flatworm eggs, and some other invertebrates. Sediments of the central/inner part of big lakes rarely contain the remains of wingless terrestrial beetles, and found fragments are strongly damaged [10]. Good burial conditions for any terrestrial insects exist in coastal parts of lakes, in the zone of plant debris accumulation (Figure 5a). Buried insect assemblages from lake shore sediments include the entire range of terrestrial and water-related taxa. The FC of water beetles and bugs is usually normal (close to 1) here, while the FC of terrestrial beetles is anomalous.

2.2. Alluvium

Fluvial deposits, such as riverbeds, old channels, and floodplains, are common in Beringia (Figure 6). With a few exceptions, insects are rarely found in the modern riverbed alluvium (alluvial deposits with crossbedding), but in the Pleistocene situation was different. There are several sites (Figure 6c) where cross-bedded sandy-gravelly alluvial deposits yielded plenty of plant debris and insect fragments. Surprisingly, the richest beetle assemblages in Beringia (Titaluk and Ikpikpuk Rivers of the North Slope of Alaska, and Keremesit, Khomus-Yuryakh and Adycha Rivers from Yakutia) were collected from this type of alluvium [8,11]. While on average, a standard 40 to 50 L sample contains 100 to 200 insect fragments, the number of fragments in the above-mentioned Beringian alluvium varied from 600 fragments per ten L sample to 8000 fragments per standard sample, with the set of species typical for the region [8].
Modern floodplain deposits commonly include layers or lenses of plant debris (Figure 5b). Similar layers were common in the Pleistocene; they yield a highly variable number of insect fragments upon wet sieving (Figure 5c,d) [10]. Probably, plant debris and dead insects were brought to the riverbed during the low water season by the river current and wind, and this material was trapped in zigzag structures at the edges of sand beaches, in their contact zone with flowing water (Figure 6a). Insect remains have been distinctly sorted there, with the hard and round heads and pronotums accumulated within gravel lenses, and flat light elytra concentrated in sand deposits and layers of plant debris. FC of all insects was anomalous. The proportion of water insects was relatively low. Ecological affiliation of insects from the assemblage is mixed without the strong dominance of either group.
Holocene floodplain deposits are often sampled in the river terraces (Figure 6d,e). Insect assemblages from the Holocene floodplain deposits are not necessarily numerous, but usually taxonomically diverse. FC may be either normal or anomalous. The proportion of water insects is often low, but riparian species are well represented. Preservation varies from poor to good.

2.3. Peatland Deposits

Peat seems to be an excellent repository of subfossil insects. However, our experience shows that this assumption is not always true. For example, a sample from the promisingly looking Holocene peat layer (Figure 1c) contained no insects. Sphagnum peat is usually insect-free; the best yield is expected from the peat of mixed origin (mosses, grasses, and leaves) and the peat that contains mineral inclusions. Bog tussocks are often exposed in Beringian sections (Figure 7), and insects can be found under the tussocks and between them. If the peat contains insects, the FC here is usually normal, preservation is good, and water and wetland taxa are well presented [12,13,14]. Sometimes organic-rich paleosols are described as “peat beds” in the field. In this case, we notice a lack of wetland taxa among plants and insects [8].

2.4. Wetland Deposits

Distinctive “grassy” deltaic deposits were observed in the delta of the Lena River (Figure 8). The deposits consist of tall stems of wetland grassy species interspersed with a very small proportion of sand. Sampling such sediment is hard work, with the result not worth the effort. Insect remains are rare in the “grassy deposits” and include mainly aquatic species, such as caddisflies [15].

2.5. Subaerial Deposits

Classic taphonomy suggests that subaerial surfaces are generally unstable in time, with a low probability of fossilization and of long-term preservation of insect remains [1]. Fossil insects are either absent or extremely rare in Pre-Quaternary aeolian deposits and paleosols, and even in the Quaternary subaerial deposits, insect remains are preserved selectively. Mid-latitudes loesses and paleosols in dry sites rarely contain any insect remains, but paleosols and archeological sites in wet settings can be rich in insects. Northern localities provide a better preservation of organic inclusions due to permafrost. In Beringia, paleosols (Figure 8) are the best source of subfossil insects [16,17]. Impressive assemblages were collected from paleosols buried under volcanic tephra [18,19,20]. Insects from buried paleosols are well preserved, FC is normal, species of true aquatic invertebrates are absent, but some water beetles that can migrate to the soil for wintering are found.
We have observed accumulation of insect remains in a modern sand dune (Figure 9d) at the Lena River [21]. A layer of plant detritus buried under the dune sand yielded insect fragments that were exhumed by wind from the soil buried nearby. Such paleosol-aeolian transfer and redeposition was probably common in the wind-prone Pleistocene landscapes of Beringia.

3. Results

The method of paleoenvironmental reconstruction that involves macroremains of insects is based on the analysis of shares of certain ecological groups [22,23,24]. Water-related insects belong to two groups: aq (aquatic) and ri (riparian and wetland). The aquatic group (Table 1) includes beetles of the following families: Gyrinidae, Haliplidae, Dytiscidae, Hydraenidae, Dryopidae, some Hydrophilidae, and a few Curculionidae; water bugs Veliidae and Corixidae (Figure 10); larvae of Ephemeroptera, Odonata, and Trichoptera (Figure 11). Aquatic beetles and bugs can migrate from water to terrain and occur in nearby soil, but their larvae reside in water permanently. So, the finds of these strictly aquatic insects are indicative of the aquatic environment, while other insects can be found outside of water bodies. For the purposes of this study, we did not include in the analysis small aquatic Diptera.
The group “ri” (Table 2) includes beetles of families Carabidae, Hydrophilidae, Staphylinidae, Scarabaeidae, Scirtidae, Heteroceridae, Elateridae, Coccinellidae, Chrysomelidae, Curculionidae, and bugs Saldidae. These insects live on wet ground, in plant litter near water bodies (Figure 12), or on riparian and wetland vegetation (Figure 13). Many riparian species can migrate to dry soils for wintering. In the cold climate, some riparian species occasionally visit dry and sunny spots to get some heat necessary for their development. Anyway, the find of “ri” group suggests that a water body existed not far from the place of burial of the insect remains.
The Beringia database (the Siberian part by [25]; the North American part prepared for publication by Kuzmina) was used to estimate the proportions between subfossil insects belonging to different ecological groups. The list of all Beringian subfossil insects includes 107,537 individuals from 1202 samples in 100 sites (Table 3).
The list of aquatic species includes 4489 specimens from 610 samples in 70 sites. The number of individuals of aquatic insects comprises only 4% of all specimens, but the group is recorded in 51% of samples and in 70% of localities.
The list of riparian species includes 5083 individuals from 656 samples in 74 sites. The proportion of the “ri” group is slightly larger than the “aq” group. The number of specimens of riparian insects comprises 5% of the total; the group is recorded in 55% of samples and in 74% of localities. We can see that both aquatic and riparian insects are less common than could be expected according to the classic taphonomic assumptions. The role of aquatic and riparian species in the subfossil assemblages is low, and many samples do not have any near-water taxa at all.
Distribution of water-related groups by geological epochs is shown in Table 3. The proportion of the “aq” group slightly increased in the Late Neogene, was low during the Pleistocene, except the warm MIS 5 interval, and then significantly increased in the Holocene. Similarly, the proportion of the “ri” group increased in the late Neogene, decreased in the Early Pleistocene, and was low in the Middle Pleistocene; it increased during the warm interval MIS 5, remaining low in the cold stages of the Late Pleistocene, and increased in the Holocene.
The clearest reaction of entomofauna on the climatic changes is reflected in the difference between the MIS5 and MIS4-2 intervals of the Late Pleistocene. Insect assemblages from cold intervals obviously yielded less water-related taxa. It must be noted that warm climatic phases of the Early-Middle Pleistocene are poorly sampled; most of the studied assemblages belong to cold phases.
Proportions of aquatic and riparian species in assemblages from different depositional environments are presented in the Table 4. Types of deposits were determined based on their diagnostic geological features.
The lowermost share of aquatic species is observed in assemblages from paleosols. Aquatic insects are usually absent in paleosol samples, but there are some exceptions. The sample OC-45 from the Old Crow site in Yukon (Table 4) is significantly different from the other three samples, while this layer was described in the field as a paleosol. In the sample, the remains of gilled aquatic taxa and caddisfly larvae were found. While the aquatic species could potentially migrate to the paleosol from the nearby water body, the presence of caddisfly larvae suggests that this paleosol was covered by water and mixed with floodplain deposits. In other samples from paleosols, riparian insects were absent or present in low numbers, except the above-described assemblage OC-45.
Peat deposits yielded a higher-than-average share of aquatic and riparian insects with a high taxonomic diversity. The proportion of aquatic species in the peat depends on the presence of open water within the peatland.

4. Discussion

The existence of the steppe-tundra or the mammoth steppe biomes in the Pleistocene was postulated based on the studies of mammal [26,27], insect [8,9,22,28], and plant [29] remains. The steppe-tundra is an extinct non-analogue landscape that served as the favorable habitat for mammals typical of tundra (e.g., caribou) and steppe (saiga, horse, bison). In the entomofauna of this period, we can notice a combination of the tundra ground beetle Pterostichus brevicornis (Kirby, 1837) and the steppe weevil Stephanocleonus eruditus Faust 1890.
A treeless aridic landscape of the north keeps water better than the southern grasslands. There are two main factors of water accumulation: the long winters that prevent full evaporation of shallow lakes and the permafrost. Ground ice (Figure 1a–c) acts as a diligent host, incorporating most of the water, excluding the water from the atmospheric cycle, and releasing the water when the ground becomes eroded due to extra heat or mechanical damage. Numerous frost cracks turn into seasonal ponds in the spring.
The steppe-tundra vegetation existed in Beringia at least since the Early Pleistocene [27]. The tundra biome was established even earlier; it is rooted in the Late Neogene boreal forest community [30]. Many species of tundra insects probably originated from the open boggy areas inside the boreal forest. Being adapted to the treeless wetlands, they could occupy wet depressions in the Pleistocene steppe-tundra as well as in the modern wet tundra.
Hydrophilic and xerophilic parts of the steppe-tundra insect community coexisted in the same climate but the temperature range of their microhabitats could be significantly different. The Mutual Climatic Range (MCR) method allows one to calculate average temperatures of January and July in the past using the modern distribution of beetle species. Application of the MCR for steppe-tundra assemblages from the lower Kolyma River showed that temperature ranges of steppe and tundra species either overlapped slightly or did not overlap at all [31]. This means that the heat balance of wet and dry sites was very different.
The proportion of thermophilus steppe species decreased during the warm intervals [8], including the Last Interglaciation (MIS5). During MIS5, some hydrophilic species became good temperature indicators. The ground beetle Dyschirius laevifasciatus Horn 1878, which currently inhabits riparian sites not much north of the Central-Western Alberta, has been found in MIS5 deposits at the Porcupine River in the northern Yukon [32]. This find validates the hypothesis that the climate of MIS5 was warmer than modern.
The relatively low proportion of the water and riparian insects in the Pleistocene subfossil assemblages could have two explanations: (1) The role of the wetland environment in the steppe-tundra was indeed low. (2) The insects were accumulated mostly in aeolian sediments and paleosols.
The origins of the Yedoma, the ice-rich silty deposit with huge ice wedges, dated by the Late Pleistocene, were long discussed by permafrost experts and paleontologists. The Yedoma (Figure 1a–c and Figure 4b,c) are widespread in both parts of Beringia. Its frozen silt contains well-preserved remains of plants and animals, including insects. Numerous insect samples come from Yedoma because of its richness; we can take a sample anywhere within the Yedoma and expect a good result. The hypotheses of the Yedoma origin are alluvial (floodplain), deltaic, proluviual, nival, aeolian, and polygenetic [33]. Despite the type of accumulation, silt particles initially result from the cryogenic weathering.
For the Eastern Beringian analogue of the Yedoma, the “Yukon muck”, the aeolian origin is widely accepted by the Western scientific community [34]. Russian scientists mostly admit the polygenetic origin of the Yedoma with the significant role of floodplain sediments in its formation [35], but recently, the Aeolian theory gained popularity there as well. One of the features of the Yedoma is the presence of tiny rootlets of grasses (Figure 4b,c). S. Gubin [36] described the layers with rootlets as “cryopedoliths”—poorly developed cryogenic soils that were affected by the repeated influx of silt on the surface.
The aeolian dust can be deposited on all surfaces, including grassy vegetation of the steppe patches and surfaces of local water bodies. Even considering that some aquatic species can migrate from water to the soil, by finding such species, we certainly know that the water existed somewhere nearby. The waterborne origin of a sediment can be validated if we find remains of freshwater crustaceans. Ostracods is a proper group for such verification; they are common in the Yedoma, well preserved, and their small size allows for detailed sampling. Comparing the insect record (after [13]) and the ostracod record (after [37]) from the Yedoma of the reference section Mamontovy Khayata (Bykovsky Peninsula at Laptev Sea coast), we can determine what layers are of a water origin.
The 40-m-high sea bluff (Figure 14) consists of frozen silt with lenses of plant detritus and gigantic ice wedges. The accumulation started more than 46 thousand years ago; the silt unit formed during the Middle Weichselian (MW) and the Late Weichselian (LW) intervals of the Late Pleistocene. The Early Holocene peat layer is situated on the top of the section, it thicknesses varies from one to three meters.
Ostracod bulk samples were taken throughout the section except some inaccessible loci. To obtain insect assemblages, large sediment samples (40–50 L) were wet sieved during three field seasons. The record is considerably detailed except the lower part, where too little sediment was exposed and hence, available for processing.
In the lower part of the section (MW1), we can see a prominent peak of the ostracod concentration and a higher-than-average proportion of aquatic insects there (Figure 14). In the upper part of the section (MW2), the ostracods almost completely disappeared while the proportion of aquatic insects slightly decreased. Ostracods appeared again in LW1, but their proportion is significantly lower than in MW1. Aquatic insects are generally sparse there. At the next stage, LW2, ostracods are absent, and the proportion of aquatic insects is low.
An unusual picture was observed in the early Holocene peat above the Yedoma (Figure 14). Aquatic insects dominated the assemblage, contributing over 50%, but only solitary ostracods were recorded. The sample was collected from the lower contact of the peat unit with the Holocene lacustrine sediment and, probably, the ostracod finds came from the buried moss, while the majority of the insect assemblage originated from both the lacustrine and peat deposits.
Even more interesting concordance and discrepancy is observed in the Yedoma part of the section. The high concentration of ostracod in MW1 points to predominantly aquatic sedimentation. In MW1, we observe a clear discrepancy—the aquatic insects peak while the ostracods are absent. Later in LW1, the curves of both groups are correlated and indicate mixed deposition. Both curves closely repeat each other in the LW2 part. According to the complete insect record [13] the terminal stage of the late Pleistocene was arid and relatively warm. It appears that the sedimentation during that stage was predominantly aeolian.
The Yedoma of the section Mamontovy Khayata does not have clear indicators of lacustrine deposition; it consists mostly of monotonous silt, sometimes with layers of allochthonous peat of various thickness that may rarely include moss. However, aquatic insects and ostracods are recorded at most sampling depths in the section. Probably, predominantly aeolian silt deposition on terrestrial surface (day surface) alternated here with the silt deposition in short-lived shallow lakes.

5. Conclusions

Generally, aquatic riparian and wetland taxa play a secondary role in the Pleistocene insect assemblages of the Beringia. Poor presentation of these groups could be explained by the limited extent of wetlands in the cryo-arid climate and by the specifics of the regional sedimentation. Occurrence of hydrophilic and xerophilic groups of insects and the proportions of each group in subfossil assemblages was closely associated with the depositional environments. Aeolian deposits and paleosols preserved a small number of aquatic insects that have hard chitin. The beetles could die far away from their burial place, with their empty exoskeleton transported later by wind or water streams to local depressions in the terrain and lake shores. Lack of dense riparian vegetation eased transportation of the insect remains from the terrain to the water. As a result, a significant part of remains of terrestrial insects ended up in alluvial and lacustrine sites.
Mixed insect assemblages could form in loess-like deposits if the dust fell into shallow short-lived lakes. Paleosols may preserve dead beetles and bugs that migrated from nearby water to the soil for wintering.
Our co-analysis of different sedimentation types and proportions of ecological groups of insects in each depositional environment attested to the predominantly aeolian origin of the specific, ice-rich Late Pleistocene deposit called Yedoma.
Proportions of hydrophilic species increased during warm climatic stages. This could be a taphonomic phenomenon associated with the distribution of shrubs and trees, as an expansion of riparian vegetation during the warm stages would prevent free transportation of terrestrial insects from the terrain to water. Climates of the warm stages were less windy; the vegetated ground produced less dust. Hence, during the wet and warm stages, the local entomofauna was reflected in subfossil assemblages more adequately than during cold phases.
The Holocene warming initiated permafrost thawing and formation of numerous thermocarst lakes. The role of wetlands dramatically increased in the Early Holocene and that is clearly reflected in subfossil insect assemblages.

Funding

The study was supported by Russian Foundation for Basic Research (RFBR) grant Number 20-04-00165.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

All material examined in this study are openly available at the facilities listed above.

Acknowledgments

The author thanks Elena Ponomarenko who carefully read and proofed the manuscript.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Sections where insects come from: (a) Kurungnakh Island in the Lena Delta, Pleistocene peat, (b) Bykovsky Peninsula (Yakutia), ice-rich section Mamontovy Khayata, (c) Undulung River (northern Yakutia), Holocene peat, (d) Kyzyl-Syr at the Viluy River, Yakutia, peat layer in the sand, (e) Bolshoy Laykhovsky Island, section Zimovie—lake sediment with ice wedge cast.
Figure 1. Sections where insects come from: (a) Kurungnakh Island in the Lena Delta, Pleistocene peat, (b) Bykovsky Peninsula (Yakutia), ice-rich section Mamontovy Khayata, (c) Undulung River (northern Yakutia), Holocene peat, (d) Kyzyl-Syr at the Viluy River, Yakutia, peat layer in the sand, (e) Bolshoy Laykhovsky Island, section Zimovie—lake sediment with ice wedge cast.
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Figure 2. Map of Beringian regions East Beringia: AC—Alaska, Central and East; BB—Bluefish River Basin and Porcupine River; ER—Eagle River, KG Klondike Gold Field, NS—North Slope of Alaska, OC—Old Crow River. West Beringia: CY Central Yakutia, KL—Kolyma Lowland, LS—Laptev Sea coast and Lena Delta, MR—Magadan Region, SC—South Chukotka, WC—West Chukotka, YL—Yana-Indigirka Lowland. Localities mentioned in the text: (1) Kurungnakh Island Lena Delta, CY 72°21′ N 126°30′ E; (2) Samoylov Island, Lena Delta, LS 72°22′ N 126°28′ E; (3) Mamontovy Khayata, Bykovsky Peninsula, LS, 71°47′ N 129°26′ E; (4) Zimovie, Bol. Lyakhovsky Island, LS, 73°20′ N, 141°21′ E; (5) Ulakhan-Sullar, Aducha River, CY, 67°41′ N, 135°44′ E; (6) Undulung River, CY 65°54′ N 126°02′ E; (7) Kyzyl-Syr, Vilyuy River, CY, 63°58′ N 123°24′ E; (8) Kharyyalakh middle Lena River, CY, 63°10′ N 129°44′ E; (9) Peschanaya Gora, middle Lena River, CY, 62°88′ N 129°81′ E; (10) Ust-Buotama, middle Lena River, CY, 61°14′ N 128°37′ E; (11) Keremesit River, YL, 70°32′ N 149°41′ E; (12) Khomus-Yuryakh River, KL, 69°59′ N, 153°35′ E; (13) Alazea River, KL, 69°20′ N, 155°00′ E; (14) Main River, SC, 64°35′ N 171°06′ E; (15) Titaluk River, NS, 69°43′ N 155°15′ W; (16) Ikpikpuk River, NS, 69°35′ N 154°55′ W; (17) Old Crow 67°53′ N 139°46′ W; (18) Porcupine River, BB, 67°29′ N 139°56′ W; (19) Hollis Mine, KG, 63°54′ N 139°26′ W; (20) Lucky Lady, KG, 63°41′ N 139°02′ W; (21) Quartz Creek, KG, 63°49′ N 139°2′ W.
Figure 2. Map of Beringian regions East Beringia: AC—Alaska, Central and East; BB—Bluefish River Basin and Porcupine River; ER—Eagle River, KG Klondike Gold Field, NS—North Slope of Alaska, OC—Old Crow River. West Beringia: CY Central Yakutia, KL—Kolyma Lowland, LS—Laptev Sea coast and Lena Delta, MR—Magadan Region, SC—South Chukotka, WC—West Chukotka, YL—Yana-Indigirka Lowland. Localities mentioned in the text: (1) Kurungnakh Island Lena Delta, CY 72°21′ N 126°30′ E; (2) Samoylov Island, Lena Delta, LS 72°22′ N 126°28′ E; (3) Mamontovy Khayata, Bykovsky Peninsula, LS, 71°47′ N 129°26′ E; (4) Zimovie, Bol. Lyakhovsky Island, LS, 73°20′ N, 141°21′ E; (5) Ulakhan-Sullar, Aducha River, CY, 67°41′ N, 135°44′ E; (6) Undulung River, CY 65°54′ N 126°02′ E; (7) Kyzyl-Syr, Vilyuy River, CY, 63°58′ N 123°24′ E; (8) Kharyyalakh middle Lena River, CY, 63°10′ N 129°44′ E; (9) Peschanaya Gora, middle Lena River, CY, 62°88′ N 129°81′ E; (10) Ust-Buotama, middle Lena River, CY, 61°14′ N 128°37′ E; (11) Keremesit River, YL, 70°32′ N 149°41′ E; (12) Khomus-Yuryakh River, KL, 69°59′ N, 153°35′ E; (13) Alazea River, KL, 69°20′ N, 155°00′ E; (14) Main River, SC, 64°35′ N 171°06′ E; (15) Titaluk River, NS, 69°43′ N 155°15′ W; (16) Ikpikpuk River, NS, 69°35′ N 154°55′ W; (17) Old Crow 67°53′ N 139°46′ W; (18) Porcupine River, BB, 67°29′ N 139°56′ W; (19) Hollis Mine, KG, 63°54′ N 139°26′ W; (20) Lucky Lady, KG, 63°41′ N 139°02′ W; (21) Quartz Creek, KG, 63°49′ N 139°2′ W.
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Figure 3. Lacustrine deposit. Sedimentation with local water invertebrates: (a) middle size tundra lakes, Old Crow Flat; (b) Ulakhan-Sullar deposit with shells.
Figure 3. Lacustrine deposit. Sedimentation with local water invertebrates: (a) middle size tundra lakes, Old Crow Flat; (b) Ulakhan-Sullar deposit with shells.
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Figure 4. Sources of insects in Yedoma deposit: (a) lake in steppe meadow near Yakutsk, (b) silt with tiny grass roots, Bykovsky Peninsula, (c) sandy silt with grass roots, Main River, Chukotka.
Figure 4. Sources of insects in Yedoma deposit: (a) lake in steppe meadow near Yakutsk, (b) silt with tiny grass roots, Bykovsky Peninsula, (c) sandy silt with grass roots, Main River, Chukotka.
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Figure 5. Grass and wood detritus: (a) plant detritus at the lake shore, Lesser Slave Lake, Canada, (b) modern alluvium with plant detritus at the Lena River, (c) section Kyzyl-Syr at the Viluy River, Yakutia with plant detritus layer at the base, (d) plant detritus in the section Old Crow River, CRH 44.
Figure 5. Grass and wood detritus: (a) plant detritus at the lake shore, Lesser Slave Lake, Canada, (b) modern alluvium with plant detritus at the Lena River, (c) section Kyzyl-Syr at the Viluy River, Yakutia with plant detritus layer at the base, (d) plant detritus in the section Old Crow River, CRH 44.
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Figure 6. River valleys and alluvial deposit: (a) Adycha, Yakutia in September 2019; riverbed is partly exposed, (b) Titaluk, North Slope of Alaska, in May 2003; water level is high, floodplain is partly flooded, (c) Pleistocene alluvial deposit at the Titaluk River with plant debris and numerous insects, (d,e) young floodplain deposit at the Lena River.
Figure 6. River valleys and alluvial deposit: (a) Adycha, Yakutia in September 2019; riverbed is partly exposed, (b) Titaluk, North Slope of Alaska, in May 2003; water level is high, floodplain is partly flooded, (c) Pleistocene alluvial deposit at the Titaluk River with plant debris and numerous insects, (d,e) young floodplain deposit at the Lena River.
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Figure 7. Boggy sediment: (a) moss and tussocks sedges in tundra, Dempster Hwy, Yukon, (b) peat tussocks, top of Mamontovy Khayata section, Bykovsky Peninsula, northern Yakutia.
Figure 7. Boggy sediment: (a) moss and tussocks sedges in tundra, Dempster Hwy, Yukon, (b) peat tussocks, top of Mamontovy Khayata section, Bykovsky Peninsula, northern Yakutia.
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Figure 8. Grass-rich sedimentation: (a) grassy edge of a water body in the tundra near Anadyr, Chukotka. (b) Grassy deposits in the Holocene section on the Samoylov Island, Lena Delta.
Figure 8. Grass-rich sedimentation: (a) grassy edge of a water body in the tundra near Anadyr, Chukotka. (b) Grassy deposits in the Holocene section on the Samoylov Island, Lena Delta.
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Figure 9. Subaerial deposit: (a) modern soil, Central Yakutia, (b) late Pleistocene paleosol, Kharyyalakh, Central Yakutia, (c) aeolian deposit with a layer of plant debris and insect fragments, (d) recent aeolian deposit with plant debris and insects, Ust-Buotama dune, Central Yakutia.
Figure 9. Subaerial deposit: (a) modern soil, Central Yakutia, (b) late Pleistocene paleosol, Kharyyalakh, Central Yakutia, (c) aeolian deposit with a layer of plant debris and insect fragments, (d) recent aeolian deposit with plant debris and insects, Ust-Buotama dune, Central Yakutia.
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Figure 10. Remains of aquatic insects who can migrate from the water: (a)—Gyrinus sp. (b)—Gyrinus natator, (c)—Gyrinus opacus, (d)—Colymbetes sp., (e)—Colymbetes dolabratus, (f)—Hygrotus sp., (g)—Agabus arcticus, (h)—A. moestus, (i)—A. serricornis, (j)—A. affinis, (k)—Ilybius sp., (l)—Hydroporus fuscipennis, (m)—Oreodytes dauricus, (n)—Haliplius immaculatus, (o)—Helophorus splendidus, (p)—H. tuberculatus, (q)—Hydrobius fuscipes, (r)—Cymbiodyta marginella, (s)—Enochrus ochropterus, (t)—Coelostoma orbiculare, (u)—Hydrochara caraboides, (v)—Ochthebius cribricollis, (w)—Limnebius sp., (x)—Bagous limosus, (y)—Eubrychius velutus, (z)—Sigara fallenoidea. (a,i,k,s,w,y,z)—Kharyyalakh, Holocene; (b,r)—Kyzyl-Syr, late Pleistocene (MIS3) peat; (c,j,l,m,u)—Peschanaya Gora: (c,j,m,t)—late Pleistocene (MIS2), (l,u)—Holocene; (d,l,z);—Ulakhan-Sullar, middle Pleistocene; (e,h,q)—Old Crow: (e,q)—middle Pleistocene, (h)—late Pleistocene; (f)—Undulung River, Holocene; (g)—Porcupine River, late Pleistocene (MIS5); (n)—Alazea River, Holocene; (o,x)—Kyzyl-Syr, late Pleistocene; (p,v)—Quartz Creek: (p)—Holocene, 26—late Pleistocene (MIS3).
Figure 10. Remains of aquatic insects who can migrate from the water: (a)—Gyrinus sp. (b)—Gyrinus natator, (c)—Gyrinus opacus, (d)—Colymbetes sp., (e)—Colymbetes dolabratus, (f)—Hygrotus sp., (g)—Agabus arcticus, (h)—A. moestus, (i)—A. serricornis, (j)—A. affinis, (k)—Ilybius sp., (l)—Hydroporus fuscipennis, (m)—Oreodytes dauricus, (n)—Haliplius immaculatus, (o)—Helophorus splendidus, (p)—H. tuberculatus, (q)—Hydrobius fuscipes, (r)—Cymbiodyta marginella, (s)—Enochrus ochropterus, (t)—Coelostoma orbiculare, (u)—Hydrochara caraboides, (v)—Ochthebius cribricollis, (w)—Limnebius sp., (x)—Bagous limosus, (y)—Eubrychius velutus, (z)—Sigara fallenoidea. (a,i,k,s,w,y,z)—Kharyyalakh, Holocene; (b,r)—Kyzyl-Syr, late Pleistocene (MIS3) peat; (c,j,l,m,u)—Peschanaya Gora: (c,j,m,t)—late Pleistocene (MIS2), (l,u)—Holocene; (d,l,z);—Ulakhan-Sullar, middle Pleistocene; (e,h,q)—Old Crow: (e,q)—middle Pleistocene, (h)—late Pleistocene; (f)—Undulung River, Holocene; (g)—Porcupine River, late Pleistocene (MIS5); (n)—Alazea River, Holocene; (o,x)—Kyzyl-Syr, late Pleistocene; (p,v)—Quartz Creek: (p)—Holocene, 26—late Pleistocene (MIS3).
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Figure 11. Remains of true aquatic insects: (a)—Odonata larvae wing pad., (bd)—plant-silt cases of Trichoptera, Brachycentridae: (b)—Brachycentrus subnubilus, (c)—Micrasema sp., (d)—Micrasema gelidum, (e) sand case of Apataniidae, (f) sand case of Molanna sp., (g,h)—Limnephilus spp.: (g) plant case, (h) sand case, (i) shell case, (j) case from shells, seeds and a beetle elytron; (a,b,h)—Kharyyalakh, Holocene, (c) Old Crow, late Pleistocene (MIS5), (d,e,f) Ulakhan-Sullar, middle Pleistocene, (g) Kyzyl-Syr, late Pleistocene (MIS3) peat, (i,j) Peschanay Gora, late Pleistocene (MIS2).
Figure 11. Remains of true aquatic insects: (a)—Odonata larvae wing pad., (bd)—plant-silt cases of Trichoptera, Brachycentridae: (b)—Brachycentrus subnubilus, (c)—Micrasema sp., (d)—Micrasema gelidum, (e) sand case of Apataniidae, (f) sand case of Molanna sp., (g,h)—Limnephilus spp.: (g) plant case, (h) sand case, (i) shell case, (j) case from shells, seeds and a beetle elytron; (a,b,h)—Kharyyalakh, Holocene, (c) Old Crow, late Pleistocene (MIS5), (d,e,f) Ulakhan-Sullar, middle Pleistocene, (g) Kyzyl-Syr, late Pleistocene (MIS3) peat, (i,j) Peschanay Gora, late Pleistocene (MIS2).
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Figure 12. Remains of insects from riparian vegetation: (a)—Cyphon variabilis; (b)—Donacia sp.; (c)—Donacia crassipes; (d)—D. splendens; (e)—D. semicuprea; (f)—Plateumaris sericea; (g)—Grypus equiseti; (h)—Notaris aethiops; (i)—N. bimaculatus, (j)—Limnobaris dolorosa; (a,b,f,j) Peschanaya Gora, Holocene, (b) Ulakhan-Sullar, late Pleistocene; (d) Kyzyl-Syr, late Pleistocene (MIS3) peat; (g)—Ust-Buotama, Holocene, (h,i(a)) Lucky Lady: (h) Holocene, (i(a)) late Pleistocene (MIS 5); (i(b)) Ulakhan-Sullar, middle Pleistocene.
Figure 12. Remains of insects from riparian vegetation: (a)—Cyphon variabilis; (b)—Donacia sp.; (c)—Donacia crassipes; (d)—D. splendens; (e)—D. semicuprea; (f)—Plateumaris sericea; (g)—Grypus equiseti; (h)—Notaris aethiops; (i)—N. bimaculatus, (j)—Limnobaris dolorosa; (a,b,f,j) Peschanaya Gora, Holocene, (b) Ulakhan-Sullar, late Pleistocene; (d) Kyzyl-Syr, late Pleistocene (MIS3) peat; (g)—Ust-Buotama, Holocene, (h,i(a)) Lucky Lady: (h) Holocene, (i(a)) late Pleistocene (MIS 5); (i(b)) Ulakhan-Sullar, middle Pleistocene.
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Figure 13. Remains of insects from plant litter, muddy and sandy water edges: (a)—Elaphrus lapponicus, (b)—E. parviceps, (c)—E. angusticollis, (d)—E. clairvillei, (e)—Nebria frigida, (f)—N. rufescens, (g)—Bembidion mutatum, (h)—B. arcticum, (i)—B. umiatense, (j)—B. lapponicum, (k)—Dyschiriodes melancholicus, (l)—Dyschirius polius, (m)—Agonum sexpunctatum, (n)—A. cupreum, (o)—A. consimile, (p)—Georyssus crenulatus, (q)—Cercyon borealis, (r)—Aegalia kamtschatica, (s)—Heterocerus fenestratus, (t)—Negastrius restrictulus, (u)—Hypnoidus bicolor, (v)—Bledius viriosus, (w)—Stenus fuscipes, (x)—S. aureoles, (y)—S. palustris, (z)—Stenus sp., (AA)—Salda littoralis, (BB)—Saldula fucicola, (CC)—Teloleuca pellucens, (DD)—Chiloxanthus stellatus. ((a),BB) Ulakhan-Sullar: (a(a)) middle Pleistocene, (BB) late Pleistocene; (b,d,h,l) Lucky Lady, (g) Hollis Mine, (e,l,n,o,t,u,v) Quartz Creek: (a(b),b,d,e,n,o,u,v)—Holocene, (g): middle Pleistocene, (c,h,i,l,u)—late Pleistocene), (f,CC) Ust-Buotama, (j,k,p,r,t(a),w,DD)—Kharyyalakh, Holocene, (m)—Kyzyl-Syr, late Pleistocene (MIS3) peat, (q) Undulung River, late Pleistocene, (x,y,z)—Batagay, late Pleistocene, (AA) Kyzyl-Syr, late Pleistocene.
Figure 13. Remains of insects from plant litter, muddy and sandy water edges: (a)—Elaphrus lapponicus, (b)—E. parviceps, (c)—E. angusticollis, (d)—E. clairvillei, (e)—Nebria frigida, (f)—N. rufescens, (g)—Bembidion mutatum, (h)—B. arcticum, (i)—B. umiatense, (j)—B. lapponicum, (k)—Dyschiriodes melancholicus, (l)—Dyschirius polius, (m)—Agonum sexpunctatum, (n)—A. cupreum, (o)—A. consimile, (p)—Georyssus crenulatus, (q)—Cercyon borealis, (r)—Aegalia kamtschatica, (s)—Heterocerus fenestratus, (t)—Negastrius restrictulus, (u)—Hypnoidus bicolor, (v)—Bledius viriosus, (w)—Stenus fuscipes, (x)—S. aureoles, (y)—S. palustris, (z)—Stenus sp., (AA)—Salda littoralis, (BB)—Saldula fucicola, (CC)—Teloleuca pellucens, (DD)—Chiloxanthus stellatus. ((a),BB) Ulakhan-Sullar: (a(a)) middle Pleistocene, (BB) late Pleistocene; (b,d,h,l) Lucky Lady, (g) Hollis Mine, (e,l,n,o,t,u,v) Quartz Creek: (a(b),b,d,e,n,o,u,v)—Holocene, (g): middle Pleistocene, (c,h,i,l,u)—late Pleistocene), (f,CC) Ust-Buotama, (j,k,p,r,t(a),w,DD)—Kharyyalakh, Holocene, (m)—Kyzyl-Syr, late Pleistocene (MIS3) peat, (q) Undulung River, late Pleistocene, (x,y,z)—Batagay, late Pleistocene, (AA) Kyzyl-Syr, late Pleistocene.
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Figure 14. Distribution of aquatic and riparian insects in the section Mamontovy Khayata, Bykovsky Peninsula, Northern-East Siberia [13] compared with the ostracod record [37].
Figure 14. Distribution of aquatic and riparian insects in the section Mamontovy Khayata, Bykovsky Peninsula, Northern-East Siberia [13] compared with the ostracod record [37].
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Table 1. List of aquatic beetles from Quaternary deposit of Beringia.
Table 1. List of aquatic beetles from Quaternary deposit of Beringia.
TaxaNumber of Region
ColeopteraSectionsSamplesIndividuals
Gyrinidae
1Gyrinus aeratus Stephens, 18351310KL
2G. confinis LeConte, 1868111AC
3G. marinus Gyllenhal, 1808111OC
4G. minutus Fabricius, 1798567AC, OC; CY, KL
5G. natator (Linnaeus, 1758)91977CY, KL, SC, WC, YL
6G. opacus C.R.Sahlberg, 1817131928OC; CY, KL, LS, YL
7G. pectoralis LeConte, 1868235BB, OC
8G. pullatus Zaitsev, 1908233BB, OC
9G. wallisi Fall, 1922111OC
10Gyrinus sp.101619AC, BB, OC; CY, KL, LS, SC
Haliplidae
11Haliplus immaculicollis Harris, 1828222BB; KL
Dytiscidae
12Agabus affinis (Paykull, 1798)222OC; CY
13A. antennatus Leech, 19395517BB, OC
14A. approximatus Fall, 1922111KG
15A. arcticus (Paykull, 1798)7812AC, BB, KG, OC; CY
16A. bifarius (Kirby, 1837)111OC
17A. clavicornis Sharp, 1882111KG
18A. colymbus Leech, 1938?111OC
19A. confinis (Gyllenhal, 1808)522OC
20A. discolor (Harris, 1828)411OC
21A. elongatus (Gyllenhal, 1826)893AC, KG, OC
22A. infuscatus Aube, 1838443BB, OC
23A. inscriptus (Crotch, 1873)111OC
24A. lapponicus (Thomson, 1867)?1153KL, LS
25A. moestus (Curtis, 1835)2396317BB, KG, NS, OC; KL, LS, SC, WC, YL
26A. semipunctatus (Kirby, 1837)2412AC, BB
27A. seriatus (Say, 1823)222OC
28A. serricornis (Paykull, 1799)3621NS, OC; CY
39A. strigulosus (Crotch, 1873)111OC
30A. thomsoni (J.Sahlberg, 1871)141965AC, KG, NS; CY, KL, LS, SC, WC, YL
31A. tristis Aube, 18382511AC, OC
32Agabus sp.58271730AC, BB, KG, LS, NS, OC; CY, KL, MR, SC, WS, YL
33Colymbetes dahuricus Aube, 18373369353AC, BB, NS, OC; KL, LS, SC, WS, YL
34C. dolabratus (Paykull, 1798)1548141AC, BB, NS, OC; KL, LS, SC, WS, YL
35C. paykulli Erichson, 1837131721CY, KL, WS, YL
36Colymbetes sp.193750AC, BB, KG, NS; CY, KL, WC
37Dytiscus circumcinctus Ahrens, 1811349CY, WC
38Dytiscus sp.678BB, KG, OC; KL, LS, YL
39Graphoderes zonatus Hoppe, 1795?113CY
40Graphoderes sp.122WC
41Hydroporus acutangulus Thomson, 1856?1045147KL, LS, SC, WC, YL
42H. appalachius Sherman, 1913223AC, BB
43H. badiellus Fall, 1923323OC
44H. dentellus Fall, 1923114OC
45H. fuscipennis Schaum, 186881240AC, BB, OC; CY, LS
46H. lapponum (Gyllenhal, 1808)111937BB, KG, NS; KL, LS, SC
47H. morio Aube, 1838122359AC, BB, KG, OC; SC
48H. polaris Fall, 192321124AC, NS
49H. puberulus LeConte, 1850?112AC
50H. punctipennis J.Sahlberg, 1880111YL
51H. rufinasus Mannerheim, 1852222BB, OC
52H. signatus Mannerheim, 1852111AC
53H. tristis (Paykull 1798)222KG; CY
54Hydroporus sp.59222538AC, BB, KG, NS, OC; CY, KL, LS, MG, SC, WS, YL
55Hygrotus sp.223KG; CY
56Ilybius erichsoni (Gemminger and Harold, 1868)1192610AC, BB, KG, OC
57I. opacus (Aube, 1837)113AY
58I. picipes (Kirby, 1837)111NS
59I. subaeneus (Erichson, 1837)111AC
60Ilybius sp.111CY
61Laccophilus sp.?111KL
62Oreodytes dauricus (Motschulsky, 1860)111CY
63O. laevis (Kirby, 1837)111NS
64O. recticollis (Fall, 1926)111NS
65O. sanmarkii (C.R.Sahlberg, 1826)112338NS; CY, KL, YL
66Rhantus notatus (Fabricius, 1781)223KL, WS
67R. frontalis (Marsham, 1802)111KG
68Dytiscidae gen. indet.101223CY, KL, LS, MR, WC, YL
Hydrophilidae
69Coelostoma orbicularis (Fabricius, 1775)?111KG
70Cymbiodyta acuminata Fall, 1924334AC, KG, OC
71C. marginella (Fabricius, 1792)112CY
72C. vindicata Fall, 1924111AC
73Cymbiodyta sp.335AC, KG, NS
74Enochrus ochropterus (Marsham, 1802)111CY
75Helophorus aspericollis Angus, 1973?122WC
76H. lacustris LeConte, 1850111AC
77H. oblongus LeConte, 1850567AC, KG, OC; WC
78H. obscurellus Poppius 19077916KL, LS, WC, YL
79H. sempervarians Angus, 1970?111NS
80H. sibiricus (Motschulsky, 1860)91115KG; KL, LS, MR, WC
81H. splendidus J.Sahlberg, 188078200445AC, BB, KG, NS, OC; CY, KL, LS, WC, YL
82H. tuberculatus (Gyllenhal, 1808)111115BB, KG; CY, KL, LS, MR, WC
83Helophorus sp.152027KG, NS, OC; CY, KL, LS, WC
84Hydrobius fuscipes (Linnaeus, 1758)77123209AC, BB, KG, OC; CY, KL, LS, MR, SC, WC, YL
85Hydrochus brevis Herbst, 1793?111KL
86Hydrochara caraboides (Linnaeus, 1758)111CY
87Laccobius sp.111520BB, OC; CY, KL, LS, WC, YL
88Hydrophilidae gen. indet.667CY, KL, LS, YL
Hydraenidae
89Hydraena sp.115KG
90Limnebius sp.111CY
91Ochthebius costatellus Reitter, 1897111OC
92O. cribricollis LeConte, 1850111BB
93O. kaszabi Janssens, 1967.71219BB, KG
94Ochthebius sp.458KG; CY, YL
Dryopidae
95Dryopidae gen. indet.111KL
Curculionidae
96Bagous limosus (Gyllenhal, 1827)345CY, LS
97B. magister LeConte, 1876115OC
98Bagous sp.555LS, YL, WC
Heteroptera
Veliidae
99Microvelia umbricola Wroblewsky, 1938112KL
Corixidae
100Sigara fallenoidea (Hungerford, 1926)112CY
101Sigara sp.112239KG, NS; CY, KL, LS, SC, YL
102Corixidae gen. indet.2214KG
Ephemeroptera
103Ephemeroptera gen. indet. (nymph)235CY
Odonata
104Odonata gen. indet. (larvae)112CY
Trichoptera
Brachycentridae
105Brachycentrus subnubilus Curtis, 18343310CY
106Micrasema gelidum McLachlan, 1876111CY
107Micrasema sp.224OC; CY
Limnephilidae
108Limnephilus sp.338CY
Molannidae
109Molanna sp.111CY
Apataniidae
110Apataniidae gen. indet.111CY
111Trichoptera gen. indet. (larvae)26105520AC, BB, NS, OC; CY, KL, LS, SC, YL
Notes: Alaska and Yukon: AC—Alaska, Central and East; BB—Bluefish River Basin and Porcupine River; KG Klondike Gold Field, NS—North Slope of Alaska, OC—Old Crow River. Northern-east Siberia: CY—Central Yakutia, KL—Kolyma Lowland, LS—Laptev Sea coast and Lena Delta, MR—Magadan Region, SC—South Chukotka, WC—West Chukotka, YL—Yana-Indigirka Lowland.
Table 2. List of riparian and wetland insects from Quaternary deposit of Beringia.
Table 2. List of riparian and wetland insects from Quaternary deposit of Beringia.
TaxaNumber of Region
ColeopteraSectionsSamplesIndividuals
Carabidae
1Agonum (Agonothorax) exaratum (Mannerheim, 1853)255AC; WC
2A. (Agonothorax) impressum (Panzer, 1796)123LS
3A. (Agonum) carbonarium Dejean, 18282612KL, LS
4A. (Agonum) sp.224KL, MR
5A. (Europhilus) consimile (Gyllenhal, 1810)132057AC, BB, KG, NS, OC; KL
6A. (Europhilus) fuliginosum (Panzer, 1809)467KL, LS
7A. (Europhilus) sexpunctatum (Linnaeus, 1758)455CY
8A. (Europhilus) simile Kirby 1837468BB, KG, OC
9A. (Europhilus) sordens Kirby, 1837?115AC
10A. (Europhilus) superioris Lindroth, 1966222AC, OC
11A. (Europhilus) thoreyi Dejean, 1828223AC, BB
12A. (Europhilus) thoreyi Dej.?111AC
13A. (Europhilus) sp.81937CY, KL, WC, YL
14A. (Olisares) affine Kirby, 1837456BB, KG, OC
15A. (Olisares) ericeti quinquepunctatum Motschulsky, 184424372AC, BB, KG, OC; KL, LS, MR, SC, WC, YL
16A. (Olisares) harrisi LeConte, 184622 AC, OC
17A. (Olisares) mutatum (Gemminger & Harold, 1868)223AC, OC
18Agonum sp.162743AC, KG; CY, KL, LS, MR, SC, WC, YL
19Amara (Curtonotus) lacustris LeConte, 1855568AY, BB, OC
20Asaphidion alaskanum Wickham 1919222NS
21A. yukonense Wickham 1919111NS
22Asaphidion sp.2512AY, CY
23Bembidion (Asioperyphus) bimaculatum (Kirby, 1837)7911NS, CY, KL, SC, WC, YL
24B. (Asioperyphus) umiatense Lindroth, 196338124321AY, BB, KG, NS, OC, CY, KL, LS, SC, WC, YL
25B. (Asioperyphus) sp.114CY
26B. (Bembidion) quadrimaculatum (Linnaeus, 1761)3515BB, KL, YL
27B. (Bembidion) sp.?345KL, YL
28B. (Bracteon) foveum Motschulsky, 1844111BB
29B. (Bracteon) lapponicum Zetterstedt, 1828334CY, LS
30B. (Bracteon) sp.222CY
31B. (Dioplocampa) transparens (Gebler, 1829)222AC, BB
32B. (Eurytrachelus) interventor Lindroth, 1963111BB
33B. (Hidromicrus) semistriatum (Haldeman, 1843)359OC
34B. (Hirmoplataphus) concolor (Kirby, 1837)111OC
35B. (Hirmoplataphus) nigrum Say, 1823226AC, OC
36B. (Hirmoplataphus) salebratum (LeConte, 1848)111BB
37B. (Leuchydrium) scudderi LeConte, 1878233BB, KG
38B. (Notaphus) coloradense Hayward, 1897122AC
39B. (Notaphus) insulatum LeConte 1852112BB
40B. (Notaphus) nigripes (Kirby, 1837)223BB, KG
41B. (Notaphus) patruele Dejean, 1831223AC, OC
42B. (Notaphus) semipunctatum Donovan, 1806111BB
43B. (Notaphus) varium (Olivier, 1795)222LS
44B. (Notaphus) sp.223KG; WC
45B. (Ocydromus) obscurellum (Motschulsky, 1845)111KG; WC
46B. (Ocydromus) sp.111KG; WC
47B. (Peryphanes) yukonum Fall, 19264813BB, OC
48B. (Peryphus) petrosum Gebler, 183381013AC; CY, KL, LS, WC, YL
49B. (Peryphus) sejunctum Casey, 1918?111AC
50B. (Peryphus) sp.122646KL, LS, SC, WC, YL
51B. (Phyla) sp.?111YL
52B. (Plataphodes) arcticum Lindroth 1963214479AC, BB, KG, NS, OC; KL, LS, WC, YL
53B. (Plataphodes) complanulum (Mannerheim, 1853)115OC
54B. (Plataphodes) compressum Lindroth 1963?111KG
55B. (Plataphodes) quadrifoveolatum Mannerheim, 1843111KG
56B. (Plataphodes) sp.445BB, OC; SC, WC
57B. (Plataphus) curtulatum Casey, 1918112OC
58B. (Plataphus) gebleri (Gebler, 1833)256LS, WC
59B. (Plataphus) hyperboraeorum Munster, 192371424BB, KG, NS; LS, WC
60B. (Plataphus) kuprianovi Mannerheim, 1843118KG
61B. (Plataphus) planatum (LeConte, 1847)111OC
62B. (Plataphus) quadrifoveolatum Mannerheim, 1843111KG
63B. (Plataphus) sp.101532KG; KL, LS, SC, YL
64B. (Pseudoperyphus) chalceum (Stephens, 1828)111BB
65B. (Semicampa) roosvelti Pic, 1902235BB, OC
66B. (Trepanedoris) acutifrons LeConte, 1879?111OC
67B. (Trepanedoris) concretum Casey, 19184911AC, BB, KG, OC
68B. (Trepanedoris) fortestriatum (Motschulsky 1845)346AC, OC
69Bembidion sp.40167711
70Blethisa inexspectata Goulet and Smetana, 1983466BB, KG; KL
71B. multipunctata (Linnaeus, 1758)91518BB, OC; KL, LS, YL
72Dyschiriodes globulosus Herbst, 17845512AC, KG
73D. melancholicus (Putzeys, 1866)43107191AC, BB, KG, NS, OC; CY, KL, LS, WC, YL
74Dyschiriodes sp.71115AC, KG; CY, LS, WC
75Dyschirius aeneolus LeConte, 1850112KG
76D. integer LeConte, 185281629BB, KG, OC
77D. setosus LeConte, 1857?111KG
78D. subarcticus Lindroth, 1961113BB, KG, OC
79Elaphrus angusticollis Putzeys, 186772046BB, KG, OC
80E. cupreus Duftschmid, 1812111KL
81E. lapponicus Gyllenhal, 1810245281BB, KG; CY, KL, LS, WC, YL
82E. parviceps Van Dyke, 1925102365AC, KG, NS, OC
83E. riparius (Linnaeus, 1758)2876142BB, OC; CY, KL, LS, SC, WC, YL
84E. splendidus Fischer von Waldheim, 182881011CY, KL, SC
85E. tuberculatus Mäklin, 1878112KG
86Elaphrus sp.132936AY, BB, KG, OC, CY, LS, WC, YL
87Nebria frigida R.F.Sahlberg, 1844132436KL, LS, MR, WC, YL
88N. gyllenhali (Strom, 1768)358KG
89N. metallica Fischer von Waldheim, 1820113KG
90N. nivalis (Paykull, 1790)111519BB, KG, OC; CY, WC, YL
91N. rufescens (Strom, 1768)222KG; CY
92N. sahlbergi Fischer von Waldheim, 1828115BB
93Nebria sp.111LS
94Opisthius richardsoni Kirby, 1837111KG
95Patrobus septentrionis Dejean, 182891015AC, BB, KG, OC; CY, WC, YL
96Patrobus sp.338AC, BB, KG, OC; CY, WC, YL
97Pelophila borealis (Paykull, 1790)151724AC, BB, OC; KL, LS, WC, YL
98Platynus mannerheimii (Dejean, 1828)123KG
99Pterostichus (Argutor) patruelis (Dejean, 1831)?1214AC
100P. (Cryobius) hudsonicus LeConte, 1863.112OC
101P. (Petrophilus) uralensis Motschulsky, 1850222KL, LS
102P. (Steropus) circulosus Lindroth, 1966?111NS
103P. (Steropus) maurusiacus Mannerheim, 1825?2415KL
104Sericoda obsoleta (Say, 1823)111OC
105S. quadripunctata (De Geer, 1774)557AC, OC, LS
106Trechus apicalis Motschulsky, 1845222CY, LS
Hydrophilidae
107Cercyon borealis Baranowski, 1985111CY
108Coelostoma orbiculare Fabricius, 17756716CY, KL, YL
109Cymbiodyta marginella (Fabricius, 1792)115OC
110Cymbiodyta sp.6713CY, KL, MR, WC, YL
111Enochrus testaceus (Fabricius, 1801)111CY
112Georissus crenulatus (Rossi, 1794)111CY
113Georissus sp.2410CY
114Megasternum posticatum (Mannerheim, 1852)115KG
Staphylinidae
115Acylophorus pronus Erichson, 1840?122ER
116Bledius confusus LeConte, 1877111OC
117B. viriosus Herman, 19831110KG
118B. zophus Herman, 1983111BB
119B. (Hesperophilus) sp.139WC
120Bledius sp.446KG; CY, LS, MR
121Boreaphilus henningianus C.R.Sahlberg, 1832112KG
122Coryphium hyperboreum (Maklin, 1880)233BB, OC
123Erichsonius nanus (Horn, 1884)?111AC
124Eucnecosum brachypterum (Gravenhorst, 1802)111AC
125Geodromicus verticalis (Say, 1831)111OC
126Holoboreaphilus nordenskioeldi (Maklin, 1878)1637188AY, KG, KL, MR, LS, WS
127Kalissus nitidus LeConte, 1874111OC
128Microedus austinianus LeConte, 1874112KG
129Olophrum boreale (Paykull, 1792)4510BB, KG, OC
130O. latum Maklin, 18531544340AC, BB, KG, NS
131O. rotundicolle (C.R.Sahlberg, 1817)112298AC, ER, KG, NS, OC
132Stenus aureolus Fauvel, 1875112CY
133S. fuscipes Gravenhorst, 1802222CY
134S. palustris Erichson, 1839112CY
135Stenus sp.50112248AC, BB, KG, NS, OC, CY, KL, LS, MR, SC, WC, YL
Scarabaeidae
136Aegialia kamtschatica Motschulsky, 186061320CY, LS, WC
137A. cylindrica (Eschscholz, 1822)3411AC
138A. lacustris LeConte, 18505627BB, KG, OC
139A. terminalis Brown, 193191025AC, BB, KG
140Aegialia sp.3311AC, KG, OC
Scirtidae
141Cyphon variabilis (Thunberg, 1787)91317AC, KG, OC, CY
142Cyphon sp.141835AC, KG; CY, KL, MR, YL
Heteroceridae
143Heterocerus fenestatus (Thunberg, 1784)111CY
144Heterocerus sp.777OC; CY, KL, LS, YL
145Lanternarius brunneus (Melsheimer, 1844)778AC, BB, KG, OC
146Lapsus tristis (Mannerheim, 1853)111KG
Elateridae
147Hypnoidus rivularius (Gyllenhal, 1827)788BB, KG; CY
148Limonius aeneoniger (DeGeer, 1774)222AC; WC
149Limonius sp.111WC
150Negastrius extricatus (Fall, 1926)111KG
151Negastrius sp.61111KG, OC; KL, MR, YL
152Oedostethus femoralis LeConte, 1853133CY
Coccinellidae
153Hippodamia arctica (Schneider, 1787)183446BB, KG, OC; CY, KL, LS, MG, SC, WC, YL
154Nephus bipunctatus (Kugelann, 1794)111CY
155Scymnus sp.92129CY, KL, LS, WC
Chrysomelidae
156Donacia cazieri Marx, 1957?111OC
157D. crassipes Fabricius, 1775122CY
158D. distincta LeConte, 1851222AC, OC
159D. hirticollis Kirby, 1837?222BB, OC
160D. porosicollis Lacordaire, 1845111OC
161D. proxima Kirby, 1837111OC
162D. semicuprea Panzer, 1796111CY
163D. splendens Jacobson, 1894222CY
164Donacia sp.222941AC, KG, OC; CY, KL, LS, WC, YL
165Hippuriphila canadensis Brown, 1942111AC
166H. mancula (LeConte, 1861) 112KG
167Hydrothassa glabra (Herbst, 1783333LS
168H. hannoverana (Fabricius, 1775)2479115KL, LS, SC, WC, YL
169Plateumaris flavipes (Kirby, 1837)3313AC, BB
170P. frosti (Schaeffer, 1935)? 111KG
171P. robusta (Schaeffer, 1920)111OC
172P. sericea (Linnaeus, 1760)111CY
173Plateumaris sp.8912AC, CY, KL, WC
174Prasocuris boreella (Schaeffer, 1928)333AC, BB, KG
175P. phellandrii (Linnaeus, 1758)222LS, WC
Curculionidae
176Auleutes epilobii (Paykull, 1800)333AC, BB, KG
177A. nebulosus (LeConte, 1876)111BB
178Erirhinus aethiops (Fabricius, 1792)2455134KG, NS, OC, CY, KL, LS, MR, SC, WC, YL
179Eubrychius velutus (Beck, 1817)225CY, KL
180Grypidius equiseti (Fabricius, 1775)163050AC, BB; CY, KL, WC, YL
181Grypus mannerheimi Faust, 1881101638CY, KL, LS, WS, YL
182Limnobaris dolorosa (Goeze, 1777)111CY
183Listronotus humilis (Gyllenhal, 1834)116OC
184Lixus paraplecticus (Linnaeus, 1758)111LS
185Notaris acridulus (Linnaeus, 1758)468CY, WC, YL
186N. bimaculatus (Fabricius, 1792)49222675AY, BB, KG, OC, CY, KL, LS, SC, WC, YL
187N. eversmanni Faust 188291767KL, LS, SC, WC, YL
188N. ochoticus Korotyaev, 198492347CY, KL, LS, WC, YL
189N. puncticollis (LeConte 1876)226BB, OC
190Notaris sp.689AC, BB, OC; WC, YL
191Pelenomus quadrituberculatus (Fabricius, 1787)333LS
192P. velaris (Gyllenhal, 1827)173777CY, KL, LS, SC, WC, YL
193Pelenomus sp.92439KL, LS, WC, YL, BB, KG
194Phytobius leucogaster (Marsham, 1802)344OC, KL
195Procas lecontei Bedel, 1879345BB, OC
196Rhinoncus castor (Fabricius, 1792)345KL
197R. jakovlevi Faust, 1893111WC
Heteroptera
Saldidae
198Chiloxanthus stellatus (Curtis, 1835)489KG, OC, CY
199Chiloxanthus sp.589AC, EA, KG, NS; KL
200Salda littoralis (Linnaeus, 1758)41520AC; LS, CY, YL
201Salda sp.81318KG; KL, LS, SC, YL
202Saldula fucicola (J.Sahlberg, 1870)111CY
203Saldula sp.5813KG, NS; SC
204Teloleuca pellucens (Fabricius, 1779)224CY, LS
205Saldidae gen. indet.6810BB, KG, OC
Notes: Alaska and Yukon: AC—Alaska, Central and East; BB—Bluefish River Basin and Porcupine River; ER—Eagle River, KG Klondike Gold Field, NS—North Slope of Alaska, OC—Old Crow River. Northern-east Siberia: CY Central Yakutia, KL—Kolyma Lowland, LS—Laptev Sea coast and Lena Delta, MR—Magadan Region, SC—South Chukotka, WC—West Chukotka, YL—Yana-Indigirka Lowland.
Table 3. Occurrence of water indicators and their percentage of assemblages in different stratigraphic units.
Table 3. Occurrence of water indicators and their percentage of assemblages in different stratigraphic units.
AgeN of Aquatic
Insects
%N of Wetland and Riparian Insects%N of All Insects
Late Neogene37511615760
Early Pleistocene237442176125
Middle Pleistocene139231715442,063
Late Pleistocene (MIS5)359647185590
Late Pleistocene (MIS4-2)153031302346,047
Holocene934131058156952
Table 4. Occurrence of water indicators and their percentage of assemblages in different sedimentation types.
Table 4. Occurrence of water indicators and their percentage of assemblages in different sedimentation types.
Region, Section, Sample, AgeRecorded aq and ri SpeciesHabitatMNIN of Insects in Sample%
aqri
Buried soil
AC, Tempest Lake, KL-03-TL-003, LPChiloxanthus sp.ri152600.2
OC, CRH 44, OC-45, MPBlethisa multipunctata (Linnaeus, 1758)ri136835
Elaphrus angusticollis Putzeys, 1867ri4
E. parviceps Van Dyke, 1925ri3
E. riparius (Linnaeus, 1758)ri2
Bembidion (Asioperyphus) umiatense Lindroth, 1963ri3
Agonum (Europhilus) simile Kirby 1837ri1
Hydroporus morio Aube, 1838aq4
Agabus confinis (Gyllenhal, 1808)aq2
Colymbetes dolabratus (Paykull, 1798)aq4
Helophorus oblongus LeConte, 1850aq1
Cyphon variabilis (Thunberg, 1787)ri1
Hippodamia arctica (Schneider, 1787)ri3
Notaris puncticollis (LeConte 1876)ri1
Trichoptera gen. indet. (larvae)aq1
KG, Lucky Lady, D-D-L-9, LP–––––––––0012500
KL, Duvanny Yar, P-1212s, LP3Bembidion (Plataphus) sp.ri19002
Notaris bimaculatus (Fabricius, 1792)ri1
Peat
OC, CRH-11, OC-8, LPGyrinus opacus C.R.Sahlberg, 1817aq11392415
Pelophila borealis (Paykull, 1790)ri5
Blethisa multipunctata (Linnaeus, 1758)ri1
Bembidion (Asioperyphus) umiatense Lindroth, 1963ri2
Agonum (Europhilus) consimile (Gyllenhal, 1810)ri2
Hydroporus morio Aube, 1838aq2
H. lapponum (Gyllenhal, 1808)aq2
Agabus bifarius (Kirby, 1837)aq1
A. erichsoni (Gemminger and Harold, 1868)aq3
A. tristis Aube, 1838aq1
A. arcticus (Paykull, 1798)aq1
A. moestus (Curtis, 1835)aq3
Colymbetes dolabratus (Paykull, 1798)aq4
Olophrum latum Maklin, 1853ri7
Helophorus splendidus J.Sahlberg, 1880aq1
Hydrobius fuscipes (Linnaeus, 1758)aq1
Cyphon variabilis (Thunberg, 1787)ri1
Hippodamia arctica (Schneider, 1787)ri3
Trichoptera gen. indet. (larvae)aq14
KG, Lucky Lady, D-L-16, HElaphrus lapponicus Gyllenhal, 1810ri268315
E. angusticollis Putzeys, 1867ri2
Dyschirius integer LeConte, 1852ri1
Dyschiriodes melancholicus (Putzeys, 1866)ri1
Patrobus septentrionis Dejean, 1828ri3
Platynus mannerheimii (Dejean, 1828)ri1
Helophorus tuberculatus Gyll.aq1
Hydrobius fuscipes (Linnaeus, 1758)aq1
LS, Kurungnakh Isl., BKh -4, HBembidion (Peryphus) petrosum Gebler, 1833ri171118
Agonum (Agonum) carbonarium Dejean, 1828ri1
Agabus moestus (Curtis, 1835)aq1
Hydroporus acutangulus Thomson, 1856?aq2
Helophorus splendidus J.Sahlberg, 1880aq1
Hydrobius fuscipes (Linnaeus, 1758)aq1
Holoboreaphilus nordenskioeldi (Maklin, 1878)ri2
Stenus sp.ri1
Hydrothassa hannoverana (Fabricius, 1775)ri1
Trichoptera gen. indet. (larvae)aq3
KG, Hollis Mine D-H-41, MPDyschiriodes melancholicus (Putzeys, 1866)ri13131112
Bembidion (Trepanedoris) concretum Casey, 1918ri1
B. (Notaphus) nigripes (Kirby, 1837)ri1
Agonum (Europhilus) consimile (Gyllenhal, 1810)ri1
Hydrobius fuscipes (Linnaeus, 1758)aq1
Grassy sediment
LS, Samoylov Isl., Sam-1-3, HDyschiriodes melancholicus (Putzeys, 1866)ri1124944
Agonum (Europhilus) fuliginosum (Panzer, 1809)ri2
Agabus thomsoni (J.Sahlberg, 1871)aq4
Holoboreaphilus nordenskioeldi (Maklin, 1878)ri1
Salda sp.ri1
Trichoptera gen. indet. (larvae)aq112
Alluvium
NS, Titaluk, T03-13, MPElaphrus parviceps Van Dyke, 1925ri138932
Bembidion (Plataphodes) arcticum Lindroth, 1963ri1
B. (Asioperyphus) sordidum (Kirby, 1837)ri1
B. (Asioperyphus) umiatense Lindroth, 1963ri2
Asaphidion alaskanum Wickham 1919ri1
Pterostichus (Steropus) circulosus Lindroth, 1963ri1
Agabus serricornis (Paykull, 1799)aq3
Hydroporus polaris Fall, 1923aq1
H. lapponum (Gyllenhal, 1808)aq5
Colymbetes sp.aq1
Helophorus splendidus J.Sahlberg, 1880aq2
KL, Alazea, AL-66-2, MPBembidion (Peryphus) sp.ri124942
Agabus moestus (Curtis, 1835)aq2
Colymbetes dolabratus (Paykull, 1798)aq1
Hydroporus sp.aq1
Dryopidae gen. indet.aq1
Hydrothassa hannoverana (Fabricius, 1775)ri1
Notaris bimaculatus (Fabricius, 1792)ri2
Trichoptera gen. indet. (larvae)aq5
KL, Alazea, AL-203-6, HGyrinus aeratus Stephens, 1835aq3623213
Blethisa multipunctata (Linnaeus, 1758)ri1
Elaphrus riparius (Linnaeus, 1758)ri1
Bembidion (Asioperyphus) umiatense Lindroth, 1963ri2
Bembidion (Plataphus) sp.ri1
Agabus lapponicus (Thomson, 1867)aq2
A. serricornis (Thomson, 1867)?aq2
A. thomsoni (J.Sahlberg, 1871)aq4
Dytiscus sp.aq1
Hydroporus fuscipennis Schaum, 1868aq1
H. lapponum (Gyllenhal, 1808)aq2
Helophorus sibiricus (Motschulsky, 1860)aq1
Hydrobius fuscipes (Linnaeus, 1758)aq4
Donacia sp.ri1
Grypidius equiseti (Fabricius, 1775)ri1
Erirhinus aethiops (Fabricius, 1792)ri1
WC, Amguema, Am-3, HBembidion (Asioperyphus) umiatense Lindroth, 1963ri5143410
B. (Plataphodes) arcticum Lindroth, 1963ri2
B. (Plataphus) hyperboraeorum Munster, 1923ri3
Bembidion sp.ri1
Agonum (Agonothorax) exaratum (Mannerheim, 1853)ri1
Agabus moestus (Curtis, 1835)aq2
Hydroporus sp.aq2
Dytiscidae gen. indet.aq1
Erirhinus aethiops (Fabricius, 1792)ri1
Notaris bimaculatus (Fabricius, 1792)ri1
Trichoptera gen. indet. (larvae)aq1
Lacustrine
KL, Alazea, AL-73-1,2, EPColymbetes dolabratus (Paykull, 1798)aq4989552
Pelenomus velaris Gyll.ri1
OC, REM-78-1, OC-53, MPElaphrus sp.ri169412
Dyschiriodes melancholicus (Putzeys, 1866)ri2
Agabus sp. aq1
Gyrinus minutus Fabricius, 1798aq1
Hydrobius fuscipes (Linnaeus, 1758)aq1
Notaris bimaculatus (Fabricius, 1792)ri5
Elaphrus parviceps Van Dyke, 1925ri1
NS, Titaluk, T03-18, HAgabus sp.aq120711
Colymbetes dolabratus (Paykull, 1798)aq2
Olophrum latum Maekl.ri1
Chiloxanthus sp.ri1
KL, Medvezhyi Isls, Medv-1-11, HGyrinus opacus C.R.Sahlberg, 1817aq1902014
Elaphrus riparius (Linnaeus, 1758)ri2
Bembidion (Asioperyphus) umiatense Lindroth, 1963ri4
Bembidion (Plataphus) sp.ri2
Agabus moestus (Curtis, 1835)aq3
Colymbetes dolabratus (Paykull, 1798)aq6
Hydroporus lapponum (Gyllenhal, 1808)aq2
Helophorus splendidus J.Sahlberg, 1880aq2
Hydrobius fuscipes (Linnaeus, 1758)aq1
Stenus sp.ri1
Notaris bimaculatus (Fabricius, 1792)ri4
Sigara sp.aq3
Yedoma
LS, Kurungnakh Isl., Bkh-02-13, LPHydrothassa glabra (Herbst, 1783)ri129711
Sigara sp.aq1
Salda sp.ri1
Trichoptera gen. indet. (larvae)aq3
YL, Yana, Tums1-11, LPBembidion (Asioperyphus) umiatense Lindroth, 1963ri211943
Helophorus splendidus J.Sahlberg, 1880aq4
Notaris sp.ri1
Sigara sp.aq1
LS, B. Lyakhovsky Isl, LR-18-9, LP–––––––––0010200
KG, Quartz Creek, D-Q-11, LP–––––––––0016100
Notes: Alaska and Yukon: AC—Alaska, Central and East; BB—Bluefish River Basin and Porcupine River; KG—Klondike Gold Field; NS—North Slope of Alaska; OC—Old Crow River, Northern-east Siberia; KL—Kolyma Lowland; LS—Laptev Sea coast and Lena Delta; MR—Magadan Region; SC—South Chukotka; WC—West Chukotka; YL—Yana-Indigirka Lowland. EP—Early Pleistocene, MP—Middle Pleistocene, LP—Late Pleistocene, H—Holocene. Samples and sites description: Tempest Lake [18], Old Crow [10] Titaluk [11] Medvezhyi Islands, Kurungnakh Island, Lyakhovsky Island, Yana, Alazea, Amguema [8] Duvanny Yar [17] Klondike Gold Field.
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Kuzmina, S. Wetlands in the Pleistocene Steppe-Tundra Landscapes of Beringia, Their Insects, and the Role of Aeolian Sedimentation. Water 2023, 15, 494. https://doi.org/10.3390/w15030494

AMA Style

Kuzmina S. Wetlands in the Pleistocene Steppe-Tundra Landscapes of Beringia, Their Insects, and the Role of Aeolian Sedimentation. Water. 2023; 15(3):494. https://doi.org/10.3390/w15030494

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Kuzmina, Svetlana. 2023. "Wetlands in the Pleistocene Steppe-Tundra Landscapes of Beringia, Their Insects, and the Role of Aeolian Sedimentation" Water 15, no. 3: 494. https://doi.org/10.3390/w15030494

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