Abstract
A lake sediment record from the north-eastern European Russian Arctic was examined using palaeolimnological methods, including subfossil chironomid and diatom analysis. The objective of this study is to disentangle environmental history of the lake and climate variability during the past 2000 years. The sediment profile was divided into two main sections following changes in the lithology, separating the limno-telmatic phase between ~2000 and 1200 cal. yr BP and the lacustrine phase between ~1200 cal. yr BP and the present. Owing to the large proportion of semi-terrestrial chironomids and poor modern analogues, a reliable chironomid-based temperature reconstruction for the limno-telmatic phase was not possible. However, the lacustrine phase showed gradually cooling climate conditions from ~1200 cal. yr BP until ~700 cal. yr BP. The increase in stream chironomids within this sediment section indicates that this period may also have had increased precipitation that caused the adjacent river to overflow, subsequently transporting chironomids to the lacustrine basin. After a short-lived warm phase at ~700 cal. yr BP, the climate again cooled, and a progressive climate warming trend was evident from the most recent sediment samples, where the biological assemblages seem to have experienced an eutrophication-like response to climate warming. The temperature reconstruction showed more similarities with the climate development in the Siberian side of the Urals than with northern Europe. This study provides a characteristic archive of arctic lake ontogeny and a valuable temperature record from a remote climate-sensitive area of northern Russia.
Introduction
Anthropogenically increased concentrations of atmospheric greenhouse gases, such as methane and carbon dioxide, has already increased air temperatures and will continue to do so in the future, especially in continental high-latitude regions (Collins et al., 2007; Miller et al., 2010; Serreze et al., 2000). Climate warming in northern regions is magnified by positive net feedbacks resulting from treeline shifts and changes in surface albedo (MacDonald, 2010; Swann et al., 2010). To better understand the ongoing climate change, proxy-based climate records that precede the observational records from remote arctic regions are required to provide a spatially exhaustive palaeoclimatological perspective on long-term climate change. Thus far, most of the palaeoclimatological and palaeoenvironmental records from the north-eastern European Russian Arctic have focused on a lower-resolution Holocene timescale (Andreev and Klimanov, 2000; Paus et al., 2003; Salonen et al., 2011), but more detailed reconstructions of late-Holocene climate trends are rare from this region. Vegetation-based reconstructions have indicated a general progressive late-Holocene cooling in the north-eastern European Russian Arctic (e.g. Salonen et al., 2011), but as a result of the relatively slow response of vegetation to climate changes (Davis, 1989), these records often fail to depict signs of the recent climate warming.
Owing to their rapid response to temperature changes and well-preserved subfossil records, chironomids (Diptera: Chironomidae) are one of the most useful palaeolimnological proxies to reconstruct past climate conditions (Brooks, 2006; Eggermont and Heiri, 2012). A previous study from northern Russia demonstrated that a chironomid-based temperature reconstruction using a Russian training set (i.e. calibration set) corresponded well with instrumental temperature data, thus indicating that subfossil chironomid assemblages in lakes in northern Russia can be used for reliable palaeotemperature reconstructions (Self et al., 2011). Although climate is known to be the predominant forcing factor on chironomids (Brooks et al., 2012), in long sedimentary records, other environmental variables than temperature can temporarily influence the chironomid community composition and hence confound the reliability of a temperature reconstruction (Nyman et al., 2008; Shala et al., 2014). Therefore, a multiproxy palaeolimnological investigation is recommended to depict variability in secondary variables. For example, diatoms (Bacillariophyta) have been shown to be important palaeolimnological indicators of water chemistry (Lotter et al., 1998; Weckström et al., 1997). Although diatoms also have palaeoclimatic indicator potential, local water chemistry determinants (e.g. lithology and marine influence) overrides the influence of climatic gradients in explaining diatom distributions in circumpolar treeline lakes of northern Russia (Laing and Smol, 2000).
In this study, a radiocarbon-dated lake sediment core from the north-eastern European Russian Arctic is examined for its subfossil insect and diatom assemblages. The aim of this study is to quantitatively reconstruct changes in past summer air temperatures over the past ~2000 years using subfossil chironomids. To depict changes in limnological conditions, diatoms are used to reconstruct changes in salinity and pH. The results of this study may be valuable in describing the late-Holocene climate variability in this remote continental high-latitude region, where the present climate change is projected to have the most significant influence.
Materials and methods
Study area
The investigated lake (unnamed) is located in the lower reaches of the Moreyu River catchment, in north-eastern European Russia (68°17′N, 59°53′E). The region is dominated by lowland tundra vegetation and is characterized by discontinuous permafrost. The shallow lake occupies a former river branch of the meandering Moreyu River, with an original inflow in the south-east. The ~2-km-long lake still has floodplain or wetland stretches related to the former river system. The northern limit is separated from the active streambed of the Moreyu River by a narrow strip of sandbars and wetlands and is situated only 1 km away from the apex of the current Moreyu Delta to the north-west. The elevation of the lake is only 2–3 m above present sea level (a.p.s.l.), and field observations suggest that it could be affected by brackish water inflow at very high tides. Partly paludified tundra uplands with shallow active layer depths (<50 cm) are found to the west and east of the lake at 5–15 m a.p.s.l. (Figure 1).

The investigated lake seen from the north-west. Inset: location of the study area in north-eastern European Russia.
The closest weather stations to the study area are Varandei, located on the Barents Coast ~95 km to the north-west, and Khoseda-Khard, corresponding to the current position of the arctic treeline ~140 km to the south. The mean July temperature (1971–2000) at Varandei is 9.3 ± 2.7°C (minimum–maximum range: 2.7–14.8°C) and at Khoseda-Khard 13.2 ± 2.3°C (minimum–maximum range: 8.1–17.8°C). By simple interpolation, mean July temperature for the study area can be estimated at ~10.5°C but displays high inter-annual variability.
Sediment sequence and radiocarbon dating
The following lake water parameters were measured during the fieldwork from lake water samples just below the water surface: conductivity 379 µS cm−1, oxygen concentration 6.7 mg L−1, turbidity 12.2 NTU and pH 7.5. The sediment sampling was performed in the northern part of the lake (water depth 2.3 m), from a rubber boat with a Russian peat corer in summer 2004. Logistical constraints related to the remoteness of the study site impeded the transport of heavier coring devices which are commonly used for this kind of lake sediment sampling. The sediment sequence consists of three separate cores that were connected via lithological horizons that represent a composite depth of ~86 cm. The bottom sediment was dark brown peat (~86–60 cm), with two dark grey brown clay intervals in between ~80 and 68 cm and a longer interval on top (~60–50 cm). A grey black clay layer (49–20 cm) occurred above it and was followed by a grey black clay layer with grey brown streaks (19–16 cm). A grey brown clay interval with black streaks (16–9 cm) was followed by a grey brown clay layer (7–0 cm) on the top sediment.
Seven samples were dated (83–86, 59–62, 43–45, 42–43, 33–34, 19–20 and 14–16 cm) using the accelerator mass spectrometry (AMS) 14C dating method. The dates from two bulk sediment samples and one sample of unknown macrofossil composition, corresponding to the upper lacustrine phase of the sequence and having reversed age estimate compared with the nearest dates, were rejected from the age–depth model (Figure 2). We ascribe these older and apparently erroneous ages to a hard-water effect in bulk lake sediments and aquatic plant macrofossils. Accepted dates are from bulk peat in the lower part of the core and selected terrestrial plant macrofossils for the top of the sequence. Compared with bulk dates from lake gyttja, radiocarbon dating of Holocene peat can be considered straightforward as this deposit consists almost entirely of locally grown plant remains (Blaauw et al., 2004). However, since there are obvious discrepancies in the chronology relative to the dated material, and there is a risk of significant changes in sedimentation rates and episodic redeposition and transport of older soils or sediments potentially as a result of flooding from the sea and the river, the results should be treated very cautiously, and the proposed chronology can only be considered as tentative. The dated samples, their material and age estimates are described in Table 1, and the relationship between sediment depth and estimated ages is illustrated in Figure 2. Radiocarbon dates were calibrated into calendar years using IntCal13 (Reimer et al., 2013) in the programme Calib (Stuiver and Reimer, 1993) Rev 7.0.4.

Radiocarbon dates from the sediment sequence. The sample marked in white was rejected as an outlier together with the two dates from bulk lake sediments (not shown in this figure).
Radiocarbon dates.
Palaeoecological analyses
Subfossil chironomid analysis was performed using standard methods (Brooks et al., 2007). Subsamples of 1–5 g wet weight were treated with 10% KOH and sieved through a 100-µm mesh. The sieved samples were transferred to a Petri dish, and the chironomid head capsules were hand-picked with fine forceps under a stereo microscope at 25× magnification. The head capsules were mounted in Euparal ventral side upwards, on microscope slides. A minimum of 50 head capsules was set as a target counting sum for each sample (Heiri and Lotter, 2001; Larocque, 2001; Quinlan and Smol, 2001). The head capsules were identified using a light microscope at 400× magnification. Identifications were based on Wiederholm (1983) and Brooks et al. (2007). In addition to chironomids, other subfossil insect remains, and oribatid mite exoskeletons were enumerated from the sediment subsamples and identified according to, for example, Heiri and Lotter (2007) and Luoto (2009a).
Diatom slide preparation followed standard methods (Battarbee et al., 2001) using the water-bath technique (Renberg, 1990). Slides were mounted using Naphrax®. The diatom accumulation rate was estimated using microsphere markers (Battarbee and Kneen, 1982). Diatom nomenclature followed Krammer and Lange-Bertalot (1986–1991).
Statistical analyses
Detrended correspondence analysis (DCA) was used as an ordination method to examine temporal changes in chironomid and diatom assemblages since the lengths of the gradient in the data sets were above 2 standard deviation (SD), thus requiring the use of a unimodal method. DCAs were carried out using CANOCO 4.5 (ter Braak and Šmilauer, 2002). The DCAs were run with square-root transformed relative taxa abundances with down-weighting of rare taxa in the programme CANOCO 4.5 (ter Braak and Šmilauer, 2002).
Chironomid-based mean July air temperature reconstruction was performed using the high-latitude Eurasian training set and a two-component weighted averaging-partial least squares (WA-PLS) model (Self et al., 2011). The model with 81 calibration sites has a bootstrapped coefficient of determination (R2) of 0.92, and a root mean squared error of prediction (RMSEP) of 1.0°C with mean and maximum biases of −0.03°C and 1.38°C, respectively. The training set can be considered suitable for the core site since its limnological features and climate characteristics are within the training set gradients. Most importantly, the temperature changes expected at the downcore site are covered by the temperature gradient of the training set (8.8–19.0°C). Furthermore, the location of the downcore site is geographically inside the training set area. All data (training set and core) were square-root transformed. Sample-specific error estimates were performed using the modern analogue technique (MAT) with 999 bootstrap iterations.
Salinity (measured as conductivity) was reconstructed using the European Diatom Database (EDDI, available online) using the combined salinity data set, which provided the best representation of the subfossil diatom assemblages. In EDDI, we used the WA inverse model, which was the most suitable in terms of its predictive performance (R2 = 0.76, RMSEP = 2.28 µS cm−1, maximum bias = 0.94 µS cm−1). pH was reconstructed using the AL:PE training set and WA inverse model (R2 = 0.78, RMSEP = 0.35, maximum bias = 0.62). The reconstructions were performed with the Ernie software (Environmental Reconstruction using the EDDI diatom database) v. 1.2 (Juggins, 2001). The diatom training sets can be considered suitable for the core sites based on their limnological features. The AL:PE training set includes sites from northern European Russia (Kola Peninsula) and has a pH range of 4.5–8.0. Although the combined salinity training set does not include sites from Russia, its conductivity gradient (40–400,000 µS cm−1) well covers the expected variability at the downcore site.
The chironomid-based temperature reconstruction was compared with chironomid DCA axis 1 scores (main direction of community variance) to verify that the changes in assemblages were driven by temperature changes (Bigler et al., 2003). This was done by examining the Pearson product–moment correlation coefficient (r) and level of statistical significance (p) between the inferred temperatures and DCA scores. Furthermore, the nearest modern analogue of the fossil samples in the temperature training set was calculated using MAT with chi-squared distance and training set and core data square-root transformed to test the suitability of the training set for the fossil data.
Results
A total of 78 chironomid taxa were identified. A total of 7 of the 54 samples contained chironomids less than the minimum count size of 50 head capsules for reliable quantitative analysis. These samples were located in the lower part of the core at sediment depths of 86, 83, 80, 74, 65, 56 and 53 cm. On average, the sediment samples between 86 and 50 cm contained 42 (5–100) head capsules, and the samples between 49 and 0 cm contained 59 (50–87) head capsules. None of the taxa occurred in all the samples, but Chironomus anthracinus type was present in 49 of the 54 samples. The most abundant taxa included Tanytarsus lugens type (mean abundance 14.4%, maximum abundance 35.1%) and C. anthracinus type (mean 12.3%, maximum 32.8%).
Of the other insect remains, mosquitoes (Culicidae) and dark-winged fungus gnats (Sciaridae) were common in the early part of the sediment profile. In addition, oribatid mites had their maximum abundances in the sediment samples located below 50 cm (Figure 3), where the sediments were composed of dark brown peat and dark grey brown clay. This section also contained a high relative proportion of semi-terrestrial chironomids (Figure 3), such as Chaetocladius dentiforceps type, Pseudosmittia, Smittia foliacea type and Limnophyes (Figure 4). Stream-associated chironomids, mostly Derotanypus (Figure 4), were common in the mid-part of the sediment profile (Figure 3), while the proportion of lacustrine chironomids, such as Psectrocladius sordidellus type, C. anthracinus type and T. lugens type (Figure 4), increased from 50 cm towards the top (Figure 3). Warm-water taxa with preference for increased nutrient conditions, such as Procladius, Stempellinella and Einfeldia pagana type, increased in the topmost sediment layers (Figure 3).

Sediment lithology, volumetric abundance of zoological macroremains and relative abundance of semi-terrestrial, riverine and lacustrine chironomids from total chironomid counts in the study lake from northern Russia.

Relative chironomid abundances in the sediment core from the study lake in northern Russia.
Chironomid DCA axis gradient lengths were 3.7 for axis 1 and 1.8 for axis 2. The first axis explained 26.1% and the second axis 6.0% of the total variance in community data. Highest axis 1 scores were assigned to samples located below 50 cm (maximum of 3.7 SD at 86 cm) indicating that these samples had distinct community composition, whereas there was only minor variability (0–0.7 SD) in the samples between 49 and 0 cm. As these lowermost samples were accumulated during a limno-telmatic phase based on the lithology and presence of terrestrial and semi-terrestrial animal remains, the DCA was also run for the lacustrine samples only (samples at 49–0 cm) to be more representative for temperature-sensitive aquatic chironomids. In this DCA, the axis 1 score increased from 49 cm until they reached maximum values (1.6–1.8 SD) between 27 and 25 cm. From there onwards, the scores gradually decreased and reached 0 SD in the surface sediment sample (Figure 6, note the reversed x-axis).
In total, 133 species were found in the subfossil diatom assemblages. The diatom abundance was very low, and a certain degree of dissolution occurred at all levels. The three lowermost samples, between 86 and 56 cm, contained only few taxa, with benthic Pinnularia lundii and Navicula pusilla being the most abundant. Pinnularia lundii reached nearly 60% at the bottommost level and then decreased rapidly and remained at <5% from about 52 cm for the rest of the sequence (Figure 5). Also, taxa such as Navicula pusilla and Diploneis interrupta had their maximum abundances below 52 cm.

Relative diatom abundances, detrended correspondence analysis (DCA) axis 1 scores and diatom-inferred salinity and pH in the sediment core from the study lake in northern Russia.
The diatom assemblages changed drastically at 52 cm and remained largely the same for the rest of the sequence. Small Fragilaroid taxa (Staurosirella var. pinnata and Staurosira construens var. venter) together with large epipelic Navicula and Pinnularia taxa dominated the diatom assemblages. Two planktonic diatoms, Aulacoseira islandica and A. subarctica, occurred at low abundance (2–3%). High-conductivity taxon Amphora lybica and several brackish Nitzschia taxa (e.g. N. levidensis) were constantly present in the upper sequence. Two Stephanodiscus species appeared in the surface sample.
Diatom-based pH and salinity (conductivity) reconstructions reflect the changes in diatom composition throughout the core sequence. Lowest pH values occurred at the limno-telmatic phase (minimum 5.5), whereas the lacustrine phase was more circumneutral (6.0–7.0). Lake conductivity shows substantial variation throughout the record between 114.8 µS cm−1 at 55 cm and 7858.5 µS cm−1 at 40 cm. High conductivity values at 48, 40 and 8 cm indicate marine influence on the lake. Conductivity shows a gradual decline towards the top of the core. Diatom DCA axis 1 sample scores exhibit overall decrease with little fluctuations throughout the core.
The chironomid-based mean July air temperature reconstruction showed large variability in the limno-telmatic sediment samples with the highest value of 15.8°C at 83 cm that was followed by a steadily decreasing trend until 68 cm, where the temperature was 9.9°C (Figure 6). The temperatures again increased towards the limno-telmatic or lacustrine sediment boundary, from where onwards a gradual cooling trend was evident with the record minimum of 8.4°C reached at 26 cm. From there onwards, the values showed a progressive increase with a temperature of 14.9°C reached at the surface sediment layer. The representation of core taxa in the temperature training set was poor in the samples located in the dark brown peat and dark grey brown clay samples during the limno-telmatic phase, as the assemblages included several taxa that are absent or rare in the training data (minimum of 60% present at 86 cm; Figure 6). In general, the representation of taxa improved upwards the sediment profile, and all the samples from 15 cm onwards had representation of >90%. Similar to the taxa representation, poorest modern analogue assemblages were found from the samples below 50 cm, where all the samples failed to reach any of the set cut-off distances (Figure 6). In all, the closest analogues were not particularly good in the upper part of the sediment profile either, but when the cut-off level was set to 10 percentile chi-squared distance, most of the samples had good analogues. With the 5 percentile cut-off, 58.1% of the lacustrine samples had good modern analogues, and with 10 percentile cut-off, 79.1% of the samples had good modern analogues. A significant correlation was found between the reconstructed temperatures and chironomid DCA axis 1 scores (r = 0.57, p < 0.001), and this relationship became even stronger when the lowermost peaty sediment layers were excluded from the examination (r = 0.75, p < 0.001).

Representation of core taxa in the training set (black bars = present, grey bars = rare and white bars = absent), closest modern analogues of the fossil assemblages in the training set (black bars = close analogue, white bars = poor analogue, black reference line = 10 percentile and grey reference line 5 percentile), detrended correspondence analysis (DCA) axis 1 scores for chironomids using all downcore samples and only lacustrine samples and chironomid-inferred mean July air temperature with bootstrapped sample-specific error estimates.
Discussion
Lake ontogeny
The radiocarbon dating results from the peat samples indicated that the sediment record covered the past ~2000 years. The lithology of the lower part of the core consisted of dark sediment material with peat layers. This part of the sediment profile, accumulated between ~2000 and 1200 cal. yr BP, was characterized by terrestrial and semi-terrestrial animal remains, such as oribatid mites, mosquitoes and dark-winged fungus gnats (Figure 3), thus indicating a limno-telmatic phase in the lake ontogeny. Subfossil oribatids, for example, are typically found in sediment samples collected from the limno-telmatic contact, while they are absent or rare from the sediments collected from the actual lake basin (Luoto, 2012). In addition, the relative proportion of semi-terrestrial (or terrestrial) chironomids was high (Figure 3) and, for example, the semi-terrestrial Chaetocladius dentiforceps type (Paasivirta, 2012) reached its maximum of 80% in the lowermost part of the sediment sequence (Figure 4).
The limno-telmatic phase had a low abundance of diatoms, and their diversity was also low (Figure 5). The diatom community consisted of circumneutral mesotrophic taxa, most of which are common for the northern Ural region (Solovieva et al., 2005, 2008). Pinnularia lundii, which tends to occur in relatively high-conductivity waters (Roberts and McMinn, 1996; WFD-UKTAG, 2014), dominated the phase (Figure 5). In addition, two other taxa which had their maximum abundances in the limno-telmatic phase, Navicula pusilla and Diploneis interrupta, are typically found in coastal brackish waters (Denis, 1994; Snoeijs and Potapova, 1995) that suggest marine influence at the time.
The upper part of the sediment core, from ~1200 cal. yr BP to the present, consisted of lacustrine clay sediments and was characterized by decreased abundance in terrestrial and semi-terrestrial mites and insects, and, for example, the terrestrial dark-winged fungus gnats disappeared permanently (Figure 3). Following the decreased number of terrestrial and semi-terrestrial taxa, the invertebrate community of the lacustrine phase changed into having a strongly increased relative abundance of aquatic chironomids (Figure 3). These aquatic chironomids included typical taxa for northern Russia with intermediate temperature optima, such as Psectrocladius (P.) sordidellus type, C. anthracinus type and T. lugens type (Self et al., 2011). The above taxa are also common in oligohumic lakes of northern Finnish Lapland (Luoto et al., 2016) indicating the diminished influence of surrounding bogs on the limnology. In addition to lacustrine chironomids, riverine chironomids were common in this phase until ~700 cal. yr BP (Figure 3). Of the stream chironomids, Derotanypus was the most abundant, and it appeared to the stratigraphy already in the mid-part of the limno-telmatic phase and began to increase towards the lacustrine phase, where it reached its maximum before 700 cal. yr BP (Figure 4). Derotanypus is rare in lake sediments (Brooks et al., 2007), but it is often found in Arctic tundra streams (Medeiros et al., 2011). Therefore, it is very likely that the lake basin had river inundations between the time period of ~1800–700 cal. yr BP as a result of increased regional precipitation causing stronger stream flow events in the Moreyu River and hence river floods transporting aquatic insects and their remains into the lake basins of the catchment. Increased effective moisture is also suggested by the basin ontogeny, as it apparently turned from a wetland into a lake (Figure 3). The hydrological dynamics of rivers emptying into the Arctic Ocean are often a spring phenomenon associated with snow melting in the catchment (Yang et al., 2004), and the subsequent flooding results in an infusion of suspended materials and nutrients that also likely affected the current study lake. The increase in the abundance of Procladius, which is common in eutrophic lakes (Brooks et al., 2001; Luoto, 2011), from ~1200 cal. yr BP onwards may be related to a gradual flood-derived nutrient enrichment of the basin. However, Procladius is also known to succeed in situations where the environmental conditions continuously change (Heiri and Lotter, 2003), such as during frequent flooding events suggesting that the changes may be more related to the dynamic environment.
The diatoms indicated increased salinity and pH for the lacustrine phase, although the conditions remained relatively stable for the whole period. While the used diatom-based training set was not specifically designed for our study region and is hence not an ideal tool for the current salinity or conductivity reconstruction, the constant presence of high-conductivity diatom Amphora lybica and several brackish Nitzschia taxa, such as N. levidensis (Sylvestre et al., 2004), clearly indicates continuous marine influence on the lake. The increase in salinity was probably related to periodic inflows of brackish water from the nearby sea coast, and the elevated pH was most likely caused by the basin ontogeny from an acidic wetland into a circumneutral lake. The presence of planktonic diatoms (Aulacoseira taxa) is typical for the mesotrophic lakes of northern Urals (Solovieva et al., 2005, 2008) suggesting increased trophic conditions.
A gradually cooling climate from ~1200 until ~700 cal. yr BP is indicated by the increases in cold-adapted chironomids (Heiri et al., 2011; Luoto, 2009b; Self et al., 2011), such as Sergentia coracina type, Protanypus and Corynocera oliveri type (Figure 4). After ~700 cal. yr BP, these taxa decreased, while more warm preferring C. anthracinus type and Procladius increased suggesting warmer climate conditions towards the present. However, from 10 cm onwards, there is a clear decrease in vegetation-associated chironomids (Brodersen et al., 2001; Brooks et al., 2007; Luoto, 2010), such as Psectrocladius sordidellus type, Cricotopus intersectus type and Paratanytarsus (Figure 4) that would suggest a decrease in aquatic macrophytes related to either colder water temperatures or increased lake level. In fact, there is a slight increase in the diatom-inferred salinity at this time that could imply flooding of brackish water into the lake basin, which could influence the lake level and survival of freshwater plants. However, it is noteworthy that the abundance of all the vegetation-associated chironomids remained very low until the present, indicating that these taxa have not recovered from the environmental stress that caused their decline. Salt water intrusions can also have direct influence on chironomids (Thienpont et al., 2015), but the present results do not show any clear evidence for the increase in salt-tolerant taxa or decline of freshwater taxa during the period of increased diatom-inferred salinity. Only the abundance of Paratanytarsus, which can tolerate salt water but is also a typical freshwater taxon (Dickson et al., 2014), increases concurrently with the diatom-based salinity reconstruction (Figures 4 and 5).
In the most recent sediment layers, there was a marked increase in chironomids that have preference to eutrophic lakes (Brooks et al., 2001; Luoto, 2011), such as Procladius, Stempellinella and Einfeldia pagana type (Figure 4). In addition to the recent changes in chironomid assemblages, there was a noticeable change in the diatom assemblages as Stephanodiscus parvus, S. minutulus and Diploneis modica increased in the surface sediment layer. Brooks et al. (2001) described a similar increase in the chironomids Procladius and Einfeldia and the diatom S. parvus along an eutrophication process in a lake in the United Kingdom that would suggest that also this study lake has experienced a recent nutrient enrichment. It is possible that the appearance of Stephanodiscus species in the surface sediment sample indicating recent enrichment of the lake is a consequence of climate warming trend in the region (Solovieva et al., 2005, 2008). However, the diatom-inferred pH indicates that the lake has become slightly more acidic during the recent times that would be against the interpretation of eutrophication, but diatoms further indicate a subtle decrease in salinity in the most recent sediment samples. Variations in temperature and salinity also affect algal blooms, for which an important condition is that temperature increases and salinity decreases. Therefore, the recent changes in the aquatic communities might result from the cumulative effects of temperature and limnology. Moreover, in freshwater environments, the effects of climate change are very often similar to those observed under eutrophication (Guilizzoni et al., 2012; Luoto et al., 2015; Schindler, 2001; Visconti et al., 2008), and hence, also in the present record, the chironomid and diatom communities appear to have experienced a eutrophication-like response to recent climate warming.
Temperature reconstruction
The chironomid-based mean July air temperature reconstruction indicated high temperatures at ~2000 cal. yr BP followed by a decreasing trend until ~1700 cal. yr BP, after which the temperatures again increased towards the end of the limno-telmatic phase. However, as the sediment section between ~2000 and 1200 cal. yr BP appears to resemble more of a wetland than a lake, the chironomid-based temperature reconstruction may not be reliable. Furthermore, owing to the large proportion of semi-terrestrial chironomids (Figure 3), these samples had poor taxa representativeness (absent or rare) in the temperature training set and, consequently, poor modern analogues (Figure 6). In addition, most of these samples contained less head capsules than the minimum set for reliable quantitative analysis, and therefore, the temperature reconstruction for the whole limno-telmatic phase is potentially unreliable. In fact, the chironomid DCA axis 1 scores for the entire sediment profile showed major changes for the limno-telmatic phase where it corresponded closely with the chironomid-inferred temperatures, whereas the changes in the lacustrine phase were very small and the relationship between the DCA scores and inferred temperatures broke apart (Figure 6). However, when the DCA was performed for the lacustrine section only, the scores again showed good correlation with the inferred temperatures, suggesting that the chironomids were responding to changes in temperature. A potential problem for the temperature reconstruction also lays in the fact that riverine chironomids occurred in the samples between ~1700 and 700 cal. yr BP (Figure 3). However, previous studies from north-eastern Europe have shown that the influence of streams and river inundations on chironomid-based temperature reconstruction is not very significant (Engels et al., 2008; Luoto, 2010; Luoto and Nevalainen, 2015). Furthermore, the most abundant stream taxon, Derotanypus, was not present in the temperature training set and, therefore, does not have any direct influence on the inferred temperatures. Nonetheless, Derotanypus is associated with very cold environments (Nazarova et al., 2015) that fit well with the current chironomid-based reconstruction, which suggests coldest temperatures during the period when Derotanypus was present (Figures 4 and 6).
A major factor influencing chironomid communities is lake-level changes (Engels and Cwynar, 2011) that could potentially compromise the reliability of the reconstruction. However, a study on boreal lakes has shown that since the inlake water depth gradient is usually directly linked with the water temperature profile, it has the same control on chironomid distribution as the air temperature gradient at large spatial gradients (Luoto et al., 2014). In addition to hydrological conditions, dissolved organic carbon (DOC) and lake humic state have significant influence on chironomid communities (Larocque et al., 2006; Luoto, 2013). In this study lake, the chironomid taxa suggesting dystrophic conditions are the same as the semi-terrestrial taxa in the lower part of the profile, whereas the chironomids of the lacustrine phase are all common in oligohumic lakes. This provides evidence that there can be serious problems with the temperature reconstruction of the semi-terrestrial phase, but the lacustrine phase is probably more reliable. The influence of salinity changes has likely had no significant effect on the temperature reconstruction since true salt water chironomids (Dickson et al., 2014) are absent from the fossil assemblages (Figure 4).
Although the lacustrine phase of the sediment profile appears to provide a valid temperature reconstruction, the comparison of the trends with other climate records is difficult owing to the uncertainties in the chronology. A tentative time frame was constructed based on the four radiocarbon dates we consider reliable (Figure 2), but we cannot exclude the possibility of changes in sedimentation rates between them or the risk of redeposition of material following flooding events from the river and the sea. Therefore, the current chronology should be regarded only as tentative.
The temperature reconstruction in the lacustrine phase showed a gradual decreasing trend towards ~700 cal. yr BP (Figure 6). This temperature decline, associated with the so-called Neoglacial cooling, has also been identified from a previous pollen-based reconstruction from Lake Kharinei and Lake Tumbulovaty in north-eastern European Russia (Salonen et al., 2011). Salonen et al. (2011) showed that this cooling began already at ~3500 cal. yr BP. Because the current temperature reconstruction resembles closely this trend, the warmer chironomid-inferred temperatures in the limno-telmatic phase of the present core data may be correct in their general trends but not in their values, since the temperature variability of 6°C within this section does not appear very realistic. The late-Holocene cooling period caused a rapid retreat of Picea treeline and triggered an abrupt initiation and activation of permafrost processes and, consequently, a major expansion of the northern Russia tundra (Salonen et al., 2011). In the present record, the gradual cooling trend was interrupted by a rapid short-lived warming episode at ~700 cal. yr BP when the temperatures increased by 2.3°C (Figure 6). This period was also reflected in the sediment lithology, where the grey brown clay was accompanied by grey brown streaks (19–16 cm). A warm phase at the same time (~600–700 cal. yr BP) has also been detected in a 1000-year-long tree-ring-based reconstruction from the eastern side of northern Urals (Briffa, 2000; Briffa et al., 1995). However, we found no clear evidence of the presence of the Medieval Climate Anomaly (MCA), a warm climate episode around 1200–800 cal. yr BP that is clearly defined from the Kola Peninsula in north-western Russia (Kremenetski et al., 2004). In common with the present reconstruction, tree-ring-based temperature reconstructions from the western Urals suggests similarly a weak MCA signal (Briffa et al., 2013) indicating that the climate in our study area resembled more Siberian conditions than those of northern Europe.
After the warm phase, the chironomid-inferred temperatures decreased close to the values that were reconstructed prior to the ~700 cal. yr BP warming (Figure 6). This period corresponds with the general timing of the ‘Little Ice Age’ (LIA) in eastern parts of Northern Europe (Luoto and Helama, 2010) and has also been previously described from tree-ring records from Siberia (Briffa, 2000). The present record also showed that the temperatures began to rapidly increase about 300 years ago, after the LIA, and continued to increase towards the present, although part of this increase may be related to chironomid response to the limnological factors unrelated to climate. In fact, there is an overestimation in the inferred temperature for the surface sample compared with the modern observed (interpolated) value (reference period: 1971–2000 = 10.5°C), but these meteorological data display very high inter-annual variability and large estimation errors. In another study from the western side of the northern Ural region, a 200-year chironomid-based temperature reconstruction from Mitrofanovskoe Lake, situated in the permafrost belt within the Bol’shezemel’skaya Tundra, indicated a rapid increase since the early 1900s (Solovieva et al., 2005), thus suggesting that the present results can be realistic. Similar to our findings, also the tree ring-based evidence from the eastern side of the northern Urals indicated a progressive climate warming from ~350 cal. yr BP (Briffa et al., 1995). Again in common with our results, Briffa et al. (1995) showed that the recent warmth was unusual, with the summer temperatures of the 20th century (1901–1990) warmer than during any similar period over the last millennium. Therefore, it appears that the general trends in summer air temperatures were similar during the examined time period on opposite sides of the Ural Mountains.
Conclusion
The lake sediment record from the north-eastern European Russian Arctic indicated that the early part of the stratigraphy accumulated during a limno-telmatic phase between ~2000 and 1200 cal. yr BP and was followed by a lacustrine phase from ~1200 cal. yr BP onwards. Diatoms suggested acidic conditions for the peaty layers of the limno-telmatic phase and the invertebrate assemblages consisted mostly of terrestrial and semi-terrestrial taxa. In the lacustrine phase, these terrestrial and semi-terrestrial invertebrates strongly decreased, and aquatic chironomids began to dominate. Diatom assemblages indicated increased salinity for the lacustrine phase, which was probably related to recurrent influxes of brackish water from the sea. Between ~1200 and 700 cal. yr BP, the relative abundance of riverine chironomids increased and suggested river floods. In the most recent sediment layers, there was a clear increase in chironomids and diatoms typical of eutrophic lakes indicating nutrient enrichment superimposed on climate warming.
The chironomid-based mean July air temperature reconstruction showed elevated temperatures at ~2000 cal. yr BP followed by a decreasing trend until ~1700 cal. yr BP. The inferred temperatures again increased towards the end of the limno-telmatic phase, but owing to the large relative abundance of semi-terrestrial chironomids and the problem of poor modern analogues, the temperature reconstruction for the limno-telmatic phase may not be reliable. However, in the lacustrine phase, which consisted of temperature-sensitive aquatic chironomids and better modern analogues in the training set, there was a gradually cooling climate trend from ~1200 until ~700 cal. yr BP. The increase in stream chironomids within this sediment section indicates that this period may also have had increased precipitation that caused the adjacent stream to overflow to the study lake. After a short-lived warm phase at ~700 cal yr PB, the climate again cooled, but a progressive climate warming was evident in the most recent sediment samples. It is potentially possible that part of this inferred temperature increase may be caused by the increase in eutrophy-preferring chironomids. However, because aquatic communities may have a eutrophication-like response to climate change, it is most likely that chironomids did respond to temperature in the whole lacustrine record and hence provide a reliable climate record for the past ~1000 years in the climatically sensitive region of the north-eastern European Russian Arctic. Apparently, the reconstructed climate development was similar in its general trends compared with a record from the opposite side of the northern Ural Mountains.
Footnotes
Funding
Fieldwork was supported by the EU FP5 GLIMPSE project (contract EVK2-CT-2002-00164) to P. Kuhry. Personal funding to T.P. Luoto was provided by the Emil Aaltonen Foundation (grant no. 160156). Constructive comments provided by the two reviewers greatly helped to improve the manuscript.
