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Increasing arctic coastal erosion rates imply a greater release of sediments and organic matter into the coastal zone. With 213 sediment samples taken around Herschel Island-Qikiqtaruk, Canadian Beaufort Sea, we aimed to gain new insights on sediment dynamics and geochemical properties of a shallow arctic nearshore zone. Spatial characteristics of nearshore sediment texture (moderately to poorly sorted silt) are dictated by hydrodynamic processes, but ice-related processes also play a role. We determined organic matter (OM) distribution and inferred the origin and quality of organic carbon by C/N ratios and stable carbon isotopes delta C-13. The carbon content was higher offshore and in sheltered areas (mean: 1.0 wt.%., S.D.: 0.9) and the C/N ratios also showed a similar spatial pattern (mean: 11.1, S.D.: 3.1), while the delta C-13 (mean: -26.4 parts per thousand VPDB, S.D.: 0.4) distribution was more complex. We compared the geochemical parameters of our study with terrestrial and marine samples from other studies using a bootstrap approach. Sediments of the current study contained 6.5 times and 1.8 times less total organic carbon than undisturbed and disturbed terrestrial sediments, respectively. Therefore, degradation of OM and separation of carbon pools take place on land and continue in the nearshore zone, where OM is leached, mineralized, or transported beyond the study area.
Variations in regional temperature have widespread implications for society, but our understanding of the amplitude and origin of long-term natural variability is insufficient for accurate regional projections. This is especially the case for terrestrial temperature variability, which is currently thought to be weak over long timescales. By performing spectral analysis on climate reconstructions, produced using sedimentary pollen records from the Northern Hemisphere over the last 8,000 years, coupled with instrumental data, we provide a comprehensive estimate of regional temperature variability from annual to millennial timescales. We show that short-term random variations are overprinted by strong ocean-driven climate variability on multi-decadal and longer timescales. This may cause substantial and potentially unpredictable regional climatic shifts in the coming century, in contrast to the relatively muted and homogeneous warming projected by climate models. Due to the marine influence, regions characterized by stable oceanic climate at sub-decadal timescales experience stronger long-term variability, and continental regions with higher sub-decadal variability show weaker long-term variability. This fundamental relationship between the timescales provides a unique insight into the emergence of a marine-driven low-frequency regime governing terrestrial climate variability and sets the basis to project the amplitude of temperature fluctuations on multi-decadal timescales and longer.
Temperature variability over land is enhanced by ocean temperature fluctuations on millennial timescales, with implications for regional-scale climate change, according to an analysis of Northern Hemisphere proxy records and observations.
How fast the Northern Hemisphere (NH) forest biome tracks strongly warming climates is largely unknown. Regional studies reveal lags between decades and millennia. Here we report a conundrum: Deglacial forest expansion in the NH extra-tropics occurs approximately 4000 years earlier in a transient MPI-ESM1.2 simulation than shown by pollen-based biome reconstructions. Shortcomings in the model and the reconstructions could both contribute to this mismatch, leaving the underlying causes unresolved. The simulated vegetation responds within decades to simulated climate changes, which agree with pollen-independent reconstructions. Thus, we can exclude climate biases as main driver for differences. Instead, the mismatch points at a multi-millennial disequilibrium of the NH forest biome to the climate signal. Therefore, the evaluation of time-slice simulations in strongly changing climates with pollen records should be critically reassessed. Our results imply that NH forests may be responding much slower to ongoing climate changes than Earth System Models predict. <br /> Deglacial forest expansion in the Northern Hemisphere poses a conundrum: Model results agree with the climate signal but are several millennia ahead of reconstructed forest dynamics. The underlying causes remain unsolved.
Pollen-based biome reconstruction on the Qinghai-Tibetan Plateau during the past 15,000 years
(2022)
Reconstruction of past vegetation change is critical for better understanding the potential impact of future global change on the fragile alpine ecosystems of the Qinghai-Tibetan Plateau (QTP). In this paper, pollen assemblages comprising 58 records from the QTP, spanning the past 15 kyrs, were collected to reconstruct biome compositions using a standard approach. Six forest biomes were identified mainly on the southeastern plateau, exhibiting a pattern of gradual expansion along the eastern margin during early to mid-Holocene times. The alpine meadow biome was separately identified based on an updated scheme, and showed notable westward expansions towards lower latitudes and higher altitudes during early Holocene times. Consistent patterns of migration could also be identified for the alpine steppe biome, which moved eastward during the late Holocene after 4 ka. As the dominant biome type, temperate steppe was distributed widely over the QTP with minor migration patterns, except for a progressive expansion to lower altitudes in the late Holocene times. The desert biome was inferred mainly as covering the northwestern plateau and the Qaidam Basin, in relatively restricted areas. The spatial distribution of the reconstructed biomes represent the large-scale vegetation gradient on the QTP. Monsoonal precipitation expressed predominant controls on the development of alpine ecosystems, while the variations in desert vegetation responded to regional moisture brought by the mid-latitude Westerlies. Temperature changes played relatively minor roles in the variations of alpine vegetation, but exerted more significant impacts on the forest biomes.
Future precipitation levels remain uncertain because climate models have struggled to reproduce observed variations in temperature-precipitation correlations. Our analyses of Holocene proxy-based temperature-precipitation correlations and hydrological sensitivities from 2,237 Northern Hemisphere extratropical pollen records reveal a significant latitudinal dependence and temporal variations among the early, middle, and late Holocene. These proxy-based variations are largely consistent with patterns obtained from transient climate simulations (TraCE21k). While high latitudes and subtropical monsoon areas show mainly stable positive correlations throughout the Holocene, the mid-latitude pattern is temporally and spatially more variable. In particular, we identified a reversal from positive to negative temperature-precipitation correlations in the eastern North American and European mid-latitudes from the early to mid-Holocene that mainly related to slowed down westerlies and a switch to moisture-limited convection under a warm climate. Our palaeoevidence of past temperature-precipitation correlation shifts identifies those regions where simulating past and future precipitation levels might be particularly challenging.
Climate change has a major impact on arctic and boreal terrestrial ecosystems as warming leads to northward treeline shifts, inducing consequences for heterotrophic organisms associated with the plant taxa. To unravel ecological dependencies, we address how long-term climatic changes have shaped the co-occurrence of plants and fungi across selected sites in Siberia. We investigated sedimentary ancient DNA from five lakes spanning the last 47,000 years, using the ITS1 marker for fungi and the chloroplast P6 loop marker for vegetation metabarcoding. We obtained 706 unique fungal operational taxonomic units (OTUs) and 243 taxa for the plants. We show higher OTU numbers in dry forest tundra as well as boreal forests compared to wet southern tundra. The most abundant fungal taxa in our dataset are Pseudeurotiaceae, Mortierella, Sordariomyceta, Exophiala, Oidiodendron, Protoventuria, Candida vartiovaarae, Pseudeurotium, Gryganskiella fimbricystis, and Tricho-sporiella cerebriformis. The overall fungal composition is explained by the plant composition as revealed by redundancy analysis. The fungal functional groups show antagonistic relationships in their climate susceptibility. The advance of woody taxa in response to past warming led to an increase in the abun-dance of mycorrhizae, lichens, and parasites, while yeast and saprotroph distribution declined. We also show co-occurrences between Salicaceae, Larix, and Alnus and their associated pathogens and detect higher mycorrhizal fungus diversity with the presence of Pinaceae. Under future warming, we can expect feedbacks between fungus composition and plant diversity changes which will affect forest advance, species diversity, and ecosystem stability in arctic regions.
Forest structure and individual tree inventories of northeastern Siberia along climatic gradients
(2022)
We compile a data set of forest surveys from expeditions to the northeast of the Russian Federation, in Krasnoyarsk Krai, the Republic of Sakha (Yakutia), and the Chukotka Autonomous Okrug (59-73 degrees N, 97-169 degrees E), performed between the years 2011 and 2021. The region is characterized by permafrost soils and forests dominated by larch (Larix gmelinii Rupr. and Larix cajanderi Mayr).
Our data set consists of a plot database describing 226 georeferenced vegetation survey plots and a tree database with information about all the trees on these plots. The tree database, consisting of two tables with the same column names, contains information on the height, species, and vitality of 40 289 trees. A subset of the trees was subject to a more detailed inventory, which recorded the stem diameter at base and at breast height, crown diameter, and height of the beginning of the crown.
We recorded heights up to 28.5 m (median 2.5 m) and stand densities up to 120 000 trees per hectare (median 1197 ha(-1)), with both values tending to be higher in the more southerly areas. Observed taxa include Larix Mill., Pinus L., Picea A. Dietr., Abies Mill., Salix L., Betula L., Populus L., Alnus Mill., and Ulmus L.
In this study, we present the forest inventory data aggregated per plot. Additionally, we connect the data with different remote sensing data products to find out how accurately forest structure can be predicted from such products. Allometries were calculated to obtain the diameter from height measurements for every species group. For Larix, the most frequent of 10 species groups, allometries depended also on the stand density, as denser stands are characterized by thinner trees, relative to height. The remote sensing products used to compare against the inventory data include climate, forest biomass, canopy height, and forest loss or disturbance. We find that the forest metrics measured in the field can only be reconstructed from the remote sensing data to a limited extent, as they depend on local properties. This illustrates the need for ground inventories like those data we present here.
The data can be used for studying the forest structure of northeastern Siberia and for the calibration and validation of remotely sensed data.
Aim: The continental-scale distribution of plant functional types, such as evergreen and summergreen needle-leaf forest, is assumed to be determined by contemporary climate. However, the distribution of summergreen needle-leaf forest of larch (Larix Mill.) differs markedly between the continents, despite relatively similar climatic conditions. The reasons for these differences are little understood. Our aim is to identify potential triggers and drivers of the current distribution patterns by comparing species' bioclimatic niches, glacial refugia and postglacial recolonization patterns.
Location: Northern hemisphere.
Taxon: Species of the genus Larix (Mill.).
Methods: We compare species distribution and dominance using species ranges and sites of dominance, as well as their occurrence on modelled permafrost extent, and active layer thickness (ALT). We compare the bioclimatic niches and calculate the niche overlap between species, using the same data in addition to modern climate data. We synthesize pollen, macrofossil and ancient DNA palaeo-evidence of past Larix occurrences of the last 60,000 years and track differences in distribution patterns through time.
Results: Bioclimatic niches show large overlaps between Asian larch species and American Larix laricina. The distribution across various degrees of permafrost extent is distinctly different for Asian L. gmelinii and L. cajanderi compared to the other species, whereas the distribution on different depths of ALT is more similar among Asian and American species. Northern glacial refugia for Larix are only present in eastern Asia and Alaska.
Main Conclusion: The dominance of summergreen larches in Asia, where evergreen conifers dominate most of the rest of the boreal forests, is dependent on the interaction of several factors which allows Asian L. gmelinii and L. cajanderi to dominate where these factors coincide. These factors include the early postglacial spread out of northern glacial refugia in the absence of competitors as well as a positive feedback mechanism between frozen ground and forest.
The biodiversity of tundra areas in northern high latitudes is threatened by invasion of forests under global warming. However, poorly understood nonlinear responses of the treeline ecotone mean the timing and extent of tundra losses are unclear, but policymakers need such information to optimize conservation efforts. Our individual-based model LAVESI, developed for the Siberian tundra-taiga ecotone, can help improve our understanding. Consequently, we simulated treeline migration trajectories until the end of the millennium, causing a loss of tundra area when advancing north. Our simulations reveal that the treeline follows climate warming with a severe, century-long time lag, which is overcompensated by infilling of stands in the long run even when temperatures cool again. Our simulations reveal that only under ambitious mitigation strategies (relative concentration pathway 2.6) will ~30% of original tundra areas remain in the north but separated into two disjunct refugia.
The SiDroForest (Siberian drone-mapped forest inventory) data collection is an attempt to remedy the scarcity of forest structure data in the circumboreal region by providing adjusted and labeled tree-level and vegetation plot-level data for machine learning and upscaling purposes. We present datasets of vegetation composition and tree and plot level forest structure for two important vegetation transition zones in Siberia, Russia; the summergreen-evergreen transition zone in Central Yakutia and the tundra-taiga transition zone in Chukotka (NE Siberia). The SiDroForest data collection consists of four datasets that contain different complementary data types that together support in-depth analyses from different perspectives of Siberian Forest plot data for multi-purpose applications. i. Dataset 1 provides unmanned aerial vehicle (UAV)-borne data products covering the vegetation plots surveyed during fieldwork (Kruse et al., 2021, ). The dataset includes structure-from-motion (SfM) point clouds and red-green-blue (RGB) and red-green-near-infrared (RGN) orthomosaics. From the orthomosaics, point-cloud products were created such as the digital elevation model (DEM), canopy height model (CHM), digital surface model (DSM) and the digital terrain model (DTM). The point-cloud products provide information on the three-dimensional (3D) structure of the forest at each plot. Dataset 2 contains spatial data in the form of point and polygon shapefiles of 872 individually labeled trees and shrubs that were recorded during fieldwork at the same vegetation plots (van Geffen et al., 2021c, ). The dataset contains information on tree height, crown diameter, and species type. These tree and shrub individually labeled point and polygon shapefiles were generated on top of the RGB UVA orthoimages. The individual tree information collected during the expedition such as tree height, crown diameter, and vitality are provided in table format. This dataset can be used to link individual information on trees to the location of the specific tree in the SfM point clouds, providing for example, opportunity to validate the extracted tree height from the first dataset. The dataset provides unique insights into the current state of individual trees and shrubs and allows for monitoring the effects of climate change on these individuals in the future. Dataset 3 contains a synthesis of 10 000 generated images and masks that have the tree crowns of two species of larch ( and ) automatically extracted from the RGB UAV images in the common objects in context (COCO) format (van Geffen et al., 2021a, ). As machine-learning algorithms need a large dataset to train on, the synthetic dataset was specifically created to be used for machine-learning algorithms to detect Siberian larch species. Larix gmeliniiLarix cajanderiDataset 4 contains Sentinel-2 (S-2) Level-2 bottom-of-atmosphere processed labeled image patches with seasonal information and annotated vegetation categories covering the vegetation plots (van Geffen et al., 2021b, ). The dataset is created with the aim of providing a small ready-to-use validation and training dataset to be used in various vegetation-related machine-learning tasks. It enhances the data collection as it allows classification of a larger area with the provided vegetation classes. The SiDroForest data collection serves a variety of user communities. <br /> The detailed vegetation cover and structure information in the first two datasets are of use for ecological applications, on one hand for summergreen and evergreen needle-leaf forests and also for tundra-taiga ecotones. Datasets 1 and 2 further support the generation and validation of land cover remote-sensing products in radar and optical remote sensing. In addition to providing information on forest structure and vegetation composition of the vegetation plots, the third and fourth datasets are prepared as training and validation data for machine-learning purposes. For example, the synthetic tree-crown dataset is generated from the raw UAV images and optimized to be used in neural networks. Furthermore, the fourth SiDroForest dataset contains S-2 labeled image patches processed to a high standard that provide training data on vegetation class categories for machine-learning classification with JavaScript Object Notation (JSON) labels provided. The SiDroForest data collection adds unique insights into remote hard-to-reach circumboreal forest regions.
Alpine ecosystems on the Tibetan Plateau are being threatened by ongoing climate warming and intensified human activities. Ecological time-series obtained from sedimentary ancient DNA (sedaDNA) are essential for understanding past ecosystem and biodiversity dynamics on the Tibetan Plateau and their responses to climate change at a high taxonomic resolution. Hitherto only few but promising studies have been published on this topic. The potential and limitations of using sedaDNA on the Tibetan Plateau are not fully understood. Here, we (i) provide updated knowledge of and a brief introduction to the suitable archives, region-specific taphonomy, state-of-the-art methodologies, and research questions of sedaDNA on the Tibetan Plateau; (ii) review published and ongoing sedaDNA studies from the Tibetan Plateau; and (iii) give some recommendations for future sedaDNA study designs. Based on the current knowledge of taphonomy, we infer that deep glacial lakes with freshwater and high clay sediment input, such as those from the southern and southeastern Tibetan Plateau, may have a high potential for sedaDNA studies. Metabarcoding (for microorganisms and plants), metagenomics (for ecosystems), and hybridization capture (for prehistoric humans) are three primary sedaDNA approaches which have been successfully applied on the Tibetan Plateau, but their power is still limited by several technical issues, such as PCR bias and incompleteness of taxonomic reference databases. Setting up high-quality and open-access regional taxonomic reference databases for the Tibetan Plateau should be given priority in the future. To conclude, the archival, taphonomic, and methodological conditions of the Tibetan Plateau are favorable for performing sedaDNA studies. More research should be encouraged to address questions about long-term ecological dynamics at ecosystem scale and to bring the paleoecology of the Tibetan Plateau into a new era.
Boreal forests of Siberia play a relevant role in the global carbon cycle. However, global warming threatens the existence of summergreen larch-dominated ecosystems, likely enabling a transition to evergreen tree taxa with deeper active layers. Complex permafrost-vegetation interactions make it uncertain whether these ecosystems could develop into a carbon source rather than continuing atmospheric carbon sequestration under global warming. Consequently, shedding light on the role of current and future active layer dynamics and the feedbacks with the apparent tree species is crucial to predict boreal forest transition dynamics and thus for aboveground forest biomass and carbon stock developments. Hence, we established a coupled model version amalgamating a one-dimensional permafrost multilayer forest land-surface model (CryoGrid) with LAVESI, an individual-based and spatially explicit forest model for larch species (Larix Mill.), extended for this study by including other relevant Siberian forest species and explicit terrain. <br /> Following parameterization, we ran simulations with the coupled version to the near future to 2030 with a mild climate-warming scenario. We focus on three regions covering a gradient of summergreen forests in the east at Spasskaya Pad, mixed summergreen-evergreen forests close to Nyurba, and the warmest area at Lake Khamra in the southeast of Yakutia, Russia. Coupled simulations were run with the newly implemented boreal forest species and compared to runs allowing only one species at a time, as well as to simulations using just LAVESI. Results reveal that the coupled version corrects for overestimation of active layer thickness (ALT) and soil moisture, and large differences in established forests are simulated. We conclude that the coupled version can simulate the complex environment of eastern Siberia by reproducing vegetation patterns, making it an excellent tool to disentangle processes driving boreal forest dynamics.
We present a chronology framework named LegacyAge 1.0 containing harmonized chronologies for 2831 pollen records (downloaded from the Neotoma Paleoecology Database and the supplementary Asian datasets) together with their age control points and metadata in machine-readable data formats.
All chronologies use the Bayesian framework implemented in Bacon version 2.5.3. Optimal parameter settings of priors (accumulation.shape, memory.strength, memory.mean, accumulation.rate, and thickness) were identified based on information in the original publication or iteratively after preliminary model inspection.
The most common control points for the chronologies are radiocarbon dates (86.1 %), calibrated by the latest calibration curves (IntCal20 and SHCal20 for the terrestrial radiocarbon dates in the Northern Hemisphere and Southern Hemisphere and Marine20 for marine materials).
The original publications were consulted when dealing with outliers and inconsistencies. Several major challenges when setting up the chronologies included the waterline issue (18.8% of records), reservoir effect (4.9 %), and sediment deposition discontinuity (4.4 %).
Finally, we numerically compare the LegacyAge 1.0 chronologies to those published in the original publications and show that the reliability of the chronologies of 95.4% of records could be improved according to our assessment.
Our chronology framework and revised chronologies provide the opportunity to make use of the ages and age uncertainties in synthesis studies of, for example, pollen-based vegetation and climate change.
The LegacyAge 1.0 dataset, including metadata, datings, harmonized chronologies, and R code used, is openaccess and available at PANGAEA (https://doi.org/10.1594/PANGAEA.933132; Li et al., 2021) and Zenodo (https://doi.org/10.5281/zenodo.5815192; Li et al., 2022), respectively.
We traced diatom composition and diversity through time using diatom-derived sedimentary ancient DNA (sedaDNA) from eastern continental slope sediments off Kamchatka (North Pacific) by applying a short, diatom-specific marker on 63 samples in a DNA metabarcoding approach. The sequences were assigned to diatoms that are common in the area and characteristic of cold water. SedaDNA allowed us to observe shifts of potential lineages from species of the genus Chaetoceros that can be related to different climatic phases, suggesting that pre-adapted ecotypes might have played a role in the long-term success of species in areas of changing environmental conditions. These sedaDNA results complement our understanding of the long-term history of diatom assemblages and their general relationship to environmental conditions of the past. Sea-ice diatoms (Pauliella taeniata [Grunow] Round & Basson, Attheya septentrionalis [ostrup] R. M. Crawford and Nitzschia frigida [Grunow]) detected during the late glacial and Younger Dryas are in agreement with previous sea-ice reconstructions. A positive correlation between pennate diatom richness and the sea-ice proxy IP25 suggests that sea ice fosters pennate diatom richness, whereas a negative correlation with June insolation and temperature points to unfavorable conditions during the Holocene. A sharp increase in proportions of freshwater diatoms at similar to 11.1 cal kyr BP implies the influence of terrestrial runoff and coincides with the loss of 42% of diatom sequence variants. We assume that reduced salinity at this time stabilized vertical stratification which limited the replenishment of nutrients in the euphotic zone.
Relative pollen productivity (RPP) estimates are fractionate values, often in relation to Poaceae, that allow vegetation cover to be estimated from pollen counts with the help of models. RPP estimates are especially used in the scientific community in Europe and China, with a few studies in North America. Here we present a comprehensive compilation of available northern hemispheric RPP studies and their results arising from 51 publications with 60 sites and 131 taxa. This compilation allows scientists to identify data gaps in need of further RPP analyses but can also aid them in finding an RPP set for their study region. We also present a taxonomically harmonised, unified RPP dataset for the Northern Hemisphere and subsets for North America (including Greenland), Europe (including arctic Russia), and China, which we generated from the available studies. The unified dataset gives the mean RPP for 55 harmonised taxa as well as fall speeds, which are necessary to reconstruct vegetation cover from pollen counts and RPP values. Data are openly available at https://doi.org/10.1594/PANGAEA.922661 (Wieczorek and Herzschuh, 2020).
Issue Despite their rather similar climatic conditions, eastern Eurasia and northern North America are largely covered by different plant functional types (deciduous or evergreen boreal forest) composed of larch or pine, spruce and fir, respectively. I propose that these deciduous and evergreen boreal forests represent alternative quasi-stable states, triggered by their different northern tree refugia that reflect the different environmental conditions experienced during the Last Glacial. Evidence This view is supported by palaeoecological and environmental evidence. Once established, Asian larch forests are likely to have stabilized through a complex vegetation-fire-permafrost soil-climate feedback system. Conclusion With respect to future forest developments, this implies that Asian larch forests are likely to be governed by long-term trajectories and are therefore largely resistant to natural climate variability on time-scales shorter than millennia. The effects of regional human impact and anthropogenic global warming might, however, cause certain stability thresholds to be crossed, meaning that irreversible transitions occur and resulting in marked consequences for ecosystem services on these human-relevant time-scales.
Although sedimentary ancient DNA (sedaDNA) has been increasingly used to study paleoecological dynamics (Schulte et al., 2020), the approach has rarely been compared with the traditional method of pollen analysis for investigating past changes in the vegetation composition and diversity of Arctic treeline areas. Here, we provide a history of latitudinal floristic composition and species diversity based on a comparison ofsedaDNA and pollen data archived in three Siberian lake sediment cores spanning the mid-Holocene to the present (7.6-0 cal ka BP), from northern typical tundra to southern open larch forest in the Omoloy region. Our results show that thesedaDNA approach identifies more plant taxa found in the local vegetation communities, while the corresponding pollen analysis mainly captures the regional vegetation development and has its limitations for plant diversity reconstruction. Measures of alpha diversity were calculated based onsedaDNA data recovered from along a tundra to forest tundra to open larch forest gradient. Across all sites,sedaDNA archives provide a complementary record of the vegetation transition within each lake's catchment, tracking a distinct latitudinal vegetation type range from larch tree/alder shrub (open larch forest site) to dwarf shrub-steppe (forest tundra) to wet sedge tundra (typical tundra site). By contrast, the pollen data reveal an open landscape, which cannot distinguish the temporal changes in compositional vegetation for the open larch forest site and forest-tundra site. IncreasingLarixpollen percentages were recorded in the forest-tundra site in the last millenium although noLarixDNA was detected, suggesting that thesedaDNA approach performs better for tracking the local establishment ofLarix. Highest species richness and diversity are found in the mid-Holocene (before 4.4 ka) at the typical tundra site with a diverse range of vegetational habitats, while lowest species richness is recorded for the forest tundra where dwarf-willow habitats dominated the lake's catchment. During the late Holocene, strong declines in species richness and diversity are found at the typical tundra site with the vegetation changing to relatively simple communities. Nevertheless, plant species richness is mostly higher than at the forest-tundra site, which shows a slightly decreasing trend. Plant species richness at the open larch forest site fluctuates through time and is higher than the other sites since around 2.5 ka. Taken together, there is no evidence to suggest that the latitudinal gradients in species diversity changes are present at a millennial scale. Additionally, a weak correlation between the principal component analysis (PCA) site scores ofsedaDNA and species richness suggests that climate may not be a direct driver of species turnover within a lake's catchment. Our data suggest thatsedaDNA and pollen have different but complementary abilities for reconstructing past vegetation and species diversity along a latitude.
Siberian larch (Larix Mill.) forests dominate vast areas of northern Russia and contribute important ecosystem services to the world. It is important to understand the past dynamics of larches in order to predict their likely response to a changing climate in the future. Sedimentary ancient DNA extracted from lake sediment cores can serve as archives to study past vegetation. However, the traditional method of studying sedimentary ancient DNA-metabarcoding-focuses on small fragments, which cannot resolve Larix to species level nor allow a detailed study of population dynamics. Here, we use shotgun sequencing and hybridization capture with long-range PCR-generated baits covering the complete Larix chloroplast genome to study Larix populations from a sediment core reaching back to 6700 years from the Taymyr region in northern Siberia. In comparison with shotgun sequencing, hybridization capture results in an increase in taxonomically classified reads by several orders of magnitude and the recovery of complete chloroplast genomes of Larix. Variation in the chloroplast reads corroborates an invasion of Larix gmelinii into the range of Larix sibirica before 6700 years ago. Since then, both species have been present at the site, although larch populations have decreased with only a few trees remaining in what was once a forested area. This study demonstrates for the first time that hybridization capture applied directly to ancient DNA of plants extracted from lake sediments can provide genome-scale information and is a viable tool for studying past genomic changes in populations of single species, irrespective of a preservation as macrofossil.
Lakes cover large parts of the climatically sensitive Arctic landscape and respond rapidly to environmental change. Arctic lakes have different origins and include the predominant thermokarst lakes, which are small, young and highly dynamic, as well as large, old and stable glacial lakes. Freshwater diatoms dominate the primary producer community in these lakes and can be used to detect biotic responses to climate and environmental change. We used specific diatom metabarcoding on sedimentary DNA, combined with next-generation sequencing and diatom morphology, to assess diatom diversity in five glacial and 15 thermokarst lakes within the easternmost expanse of the Siberian treeline ecotone in Chukotka, Russia. We obtained 163 verified diatom sequence types and identified 176 diatom species morphologically. Although there were large differences in taxonomic assignment using the two approaches, they showed similar high abundances and diversity of Fragilariceae and Aulacoseiraceae. In particular, the genetic approach detected hidden within-lake variations of fragilarioids in glacial lakes and dominance of centric Aulacoseira species, whereas Lindavia ocellata was predominant using morphology. In thermokarst lakes, sequence types and valve counts also detected high diversity of Fragilariaceae, which followed the vegetation gradient along the treeline. Ordination analyses of the genetic data from glacial and thermokarst lakes suggest that concentrations of sulfate (SO42-), an indicator of the activity of sulfate-reducing microbes under anoxic conditions, and bicarbonate (HCO3-), which relates to surrounding vegetation, have a significant influence on diatom community composition. For thermokarst lakes, we also identified lake depth as an important variable, but SO42- best explains diatom diversity derived from genetic data, whereas HCO3- best explains the data from valve counts. Higher diatom diversity was detected in glacial lakes, most likely related to greater lake age and different edaphic settings, which gave rise to diversification and endemism. In contrast, small, dynamic thermokarst lakes are inhabited by stress-tolerant fragilarioids and are related to different vegetation types along the treeline ecotone. Our study demonstrated that genetic investigations of lake sediments can be used to interpret climate and environmental responses of diatoms. It also showed how lake type affects diatom diversity, and that such genetic analyses can be used to track diatom community changes under ongoing warming in the Arctic.
The occurrence of refugia beyond the arctic treeline and genetic adaptation therein play a crucial role of largely unknown effect size. While refugia have potential for rapidly colonizing the tundra under global warming, the taxa may be maladapted to the new environmental conditions. Understanding the genetic composition and age of refugia is thus crucial for predicting any migration response.
Here, we genotype 194 larch individuals from an similar to 1.8 km(2)area in northcentral Siberia on the southern Taimyr Peninsula by applying an assay of 16 nuclear microsatellite markers. For estimating the age of clonal individuals, we counted tree rings at sections along branches to establish a lateral growth rate that was then combined with geographic distance.
Findings reveal that the predominant reproduction type is clonal (58.76%) by short distance spreading of ramets. One outlier of clones 1 km apart could have been dispersed by reindeer. In clonal groups and within individuals, we find that somatic mutations accumulate with geographic distance. Clonal groups of two or more individuals are observed. Clonal age estimates regularly suggest individuals as old as 2,200 years, which coincides with a major environmental change that forced a treeline retreat in the region.
We conclude that individuals with clonal growth mode were naturally selected as it lowers the likely risk of extinction under a harsh environment. We discuss this legacy from the past that might now be a maladaptation and hinder expansion under currently strongly increasing temperatures.
The Fram Strait is an area with a relatively low and irregular distribution of diatom microfossils in surface sediments, and thus microfossil records are scarce, rarely exceed the Holocene, and contain sparse information about past richness and taxonomic composition. These attributes make the Fram Strait an ideal study site to test the utility of sedimentary ancient DNA (sedaDNA) metabarcoding. Amplifying a short, partial rbcL marker from samples of sediment core MSM05/5-712-2 resulted in 95.7% of our sequences being assigned to diatoms across 18 different families, with 38.6% of them being resolved to species and 25.8% to genus level. Independent replicates show a high similarity of PCR products, especially in the oldest samples. Diatom sedaDNA richness is highest in the Late Weichselian and lowest in Mid- and Late Holocene samples. Taxonomic composition is dominated by cold-water and sea-ice-associated diatoms and suggests several reorganisations - after the Last Glacial Maximum, after the Younger Dryas, and after the Early and after the Mid-Holocene. Different sequences assigned to, amongst others, Chaetoceros socialis indicate the detectability of intra-specific diversity using sedaDNA. We detect no clear pattern between our diatom sedaDNA record and the previously published IP25 record of this core, although proportions of pennate diatoms increase with higher IP25 concentrations and proportions of Nitzschia cf. frigida exceeding 2% of the assemblage point towards past sea-ice presence.
The Fram Strait is an area with a relatively low and irregular distribution of diatom microfossils in surface sediments, and thus microfossil records are scarce, rarely exceed the Holocene, and contain sparse information about past richness and taxonomic composition. These attributes make the Fram Strait an ideal study site to test the utility of sedimentary ancient DNA (sedaDNA) metabarcoding. Amplifying a short, partial rbcL marker from samples of sediment core MSM05/5-712-2 resulted in 95.7 % of our sequences being assigned to diatoms across 18 different families, with 38.6 % of them being resolved to species and 25.8 % to genus level. Independent replicates show a high similarity of PCR products, especially in the oldest samples. Diatom sedaDNA richness is highest in the Late Weichselian and lowest in Mid- and Late Holocene samples. Taxonomic composition is dominated by cold-water and sea-ice-associated diatoms and suggests several reorganisations – after the Last Glacial Maximum, after the Younger Dryas, and after the Early and after the Mid-Holocene. Different sequences assigned to, amongst others, Chaetoceros socialis indicate the detectability of intra-specific diversity using sedaDNA. We detect no clear pattern between our diatom sedaDNA record and the previously published IP25 record of this core, although proportions of pennate diatoms increase with higher IP25 concentrations and proportions of Nitzschia cf. frigida exceeding 2 % of the assemblage point towards past sea-ice presence.
The Fram Strait is an area with a relatively low and irregular distribution of diatom microfossils in surface sediments, and thus microfossil records are scarce, rarely exceed the Holocene, and contain sparse information about past richness and taxonomic composition. These attributes make the Fram Strait an ideal study site to test the utility of sedimentary ancient DNA (sedaDNA) metabarcoding. Amplifying a short, partial rbcL marker from samples of sediment core MSM05/5-712-2 resulted in 95.7 % of our sequences being assigned to diatoms across 18 different families, with 38.6 % of them being resolved to species and 25.8 % to genus level. Independent replicates show a high similarity of PCR products, especially in the oldest samples. Diatom sedaDNA richness is highest in the Late Weichselian and lowest in Mid- and Late Holocene samples. Taxonomic composition is dominated by cold-water and sea-ice-associated diatoms and suggests several reorganisations – after the Last Glacial Maximum, after the Younger Dryas, and after the Early and after the Mid-Holocene. Different sequences assigned to, amongst others, Chaetoceros socialis indicate the detectability of intra-specific diversity using sedaDNA. We detect no clear pattern between our diatom sedaDNA record and the previously published IP25 record of this core, although proportions of pennate diatoms increase with higher IP25 concentrations and proportions of Nitzschia cf. frigida exceeding 2 % of the assemblage point towards past sea-ice presence.
Aim This study investigates taxonomic and phylogenetic diversity in diatom genera to evaluate assembly rules for eukaryotic microbes across the Siberian tree line. We first analysed how phylogenetic distance relates to taxonomic richness and turnover. Second, we used relatedness indices to evaluate if environmental filtering or competition influences the assemblies in space and through time. Third, we used distance-based ordination to test which environmental variables shape diatom turnover. Location Yakutia and Taymyria, Russia: we sampled 78 surface sediments and a sediment core, extending to 7,000 years before present, to capture the forest-tundra transition in space and time respectively. Taxon Arctic freshwater diatoms. Methods We applied metabarcoding to retrieve diatom diversity from surface and core sedimentary DNA. The taxonomic assignment binned sequence types (lineages) into genera and created taxonomic (abundance of lineages within different genera) and phylogenetic datasets (phylogenetic distances of lineages within different genera). Results Contrary to our expectations, we find a unimodal relationship between phylogenetic distance and richness in diatom genera. We discern a positive relationship between phylogenetic distance and taxonomic turnover in spatially and temporally distributed diatom genera. Furthermore, we reveal positive relatedness indices in diatom genera across the spatial environmental gradient and predominantly in time slices at a single location, with very few exceptions assuming effects of competition. Distance-based ordination of taxonomic and phylogenetic turnover indicates that lake environment variables, like HCO3- and water depth, largely explain diatom turnover. Main conclusion Phylogenetic and abiotic assembly rules are important in understanding the regional assembly of diatom genera across lakes in the Siberian tree line ecotone. Using a space-time approach we are able to exclude the influence of geography and elucidate that lake environmental variables primarily shape the assemblies. We conclude that some diatom genera have greater capabilities to adapt to environmental changes, whereas others will be putatively replaced or lost due to the displacement of the Arctic tundra biome under recent global warming.
Pollen records from Siberia are mostly absent in global or Northern Hemisphere synthesis works. Here we present a taxonomically harmonized and temporally standardized pollen dataset that was synthesized using 173 palynological records from Siberia and adjacent areas (northeastern Asia, 42-75 degrees N, 50-180 degrees E). Pollen data were taxonomically harmonized, i.e. the original 437 taxa were assigned to 106 combined pollen taxa. Age-depth models for all records were revised by applying a constant Bayesian age-depth modelling routine. The pollen dataset is available as count data and percentage data in a table format (taxa vs. samples), with age information for each sample. The dataset has relatively few sites covering the last glacial period between 40 and 11.5 ka (calibrated thousands of years before 1950 CE) particularly from the central and western part of the study area. In the Holocene period, the dataset has many sites from most of the area, with the exception of the central part of Siberia. Of the 173 pollen records, 81 % of pollen counts were downloaded from open databases (GPD, EPD, PANGAEA) and 10 % were contributions by the original data gatherers, while a few were digitized from publications. Most of the pollen records originate from peatlands (48 %) and lake sediments (33 %). Most of the records (83 %) have >= 3 dates, allowing the establishment of reliable chronologies. The dataset can be used for various purposes, including pollen data mapping (example maps for Larix at selected time slices are shown) as well as quantitative climate and vegetation reconstructions. The datasets for pollen counts and pollen percentages are available at https://doi.org/10.1594/PANGAEA.898616 (Cao et al., 2019a), also including the site information, data source, original publication, dating data, and the plant functional type for each pollen taxa.
Pollen records from Siberia are mostly absent in global or Northern Hemisphere synthesis works. Here we present a taxonomically harmonized and temporally standardized pollen dataset that was synthesized using 173 palynological records from Siberia and adjacent areas (northeastern Asia, 42-75 degrees N, 50-180 degrees E). Pollen data were taxonomically harmonized, i.e. the original 437 taxa were assigned to 106 combined pollen taxa. Age-depth models for all records were revised by applying a constant Bayesian age-depth modelling routine. The pollen dataset is available as count data and percentage data in a table format (taxa vs. samples), with age information for each sample. The dataset has relatively few sites covering the last glacial period between 40 and 11.5 ka (calibrated thousands of years before 1950 CE) particularly from the central and western part of the study area. In the Holocene period, the dataset has many sites from most of the area, with the exception of the central part of Siberia. Of the 173 pollen records, 81 % of pollen counts were downloaded from open databases (GPD, EPD, PANGAEA) and 10 % were contributions by the original data gatherers, while a few were digitized from publications. Most of the pollen records originate from peatlands (48 %) and lake sediments (33 %). Most of the records (83 %) have >= 3 dates, allowing the establishment of reliable chronologies. The dataset can be used for various purposes, including pollen data mapping (example maps for Larix at selected time slices are shown) as well as quantitative climate and vegetation reconstructions. The datasets for pollen counts and pollen percentages are available at https://doi.org/10.1594/PANGAEA.898616 (Cao et al., 2019a), also including the site information, data source, original publication, dating data, and the plant functional type for each pollen taxa.
Aim This study investigates taxonomic and phylogenetic diversity in diatom genera to evaluate assembly rules for eukaryotic microbes across the Siberian tree line. We first analysed how phylogenetic distance relates to taxonomic richness and turnover. Second, we used relatedness indices to evaluate if environmental filtering or competition influences the assemblies in space and through time. Third, we used distance-based ordination to test which environmental variables shape diatom turnover. Location Yakutia and Taymyria, Russia: we sampled 78 surface sediments and a sediment core, extending to 7,000 years before present, to capture the forest-tundra transition in space and time respectively. Taxon Arctic freshwater diatoms. Methods We applied metabarcoding to retrieve diatom diversity from surface and core sedimentary DNA. The taxonomic assignment binned sequence types (lineages) into genera and created taxonomic (abundance of lineages within different genera) and phylogenetic datasets (phylogenetic distances of lineages within different genera). Results Contrary to our expectations, we find a unimodal relationship between phylogenetic distance and richness in diatom genera. We discern a positive relationship between phylogenetic distance and taxonomic turnover in spatially and temporally distributed diatom genera. Furthermore, we reveal positive relatedness indices in diatom genera across the spatial environmental gradient and predominantly in time slices at a single location, with very few exceptions assuming effects of competition. Distance-based ordination of taxonomic and phylogenetic turnover indicates that lake environment variables, like HCO3- and water depth, largely explain diatom turnover. Main conclusion Phylogenetic and abiotic assembly rules are important in understanding the regional assembly of diatom genera across lakes in the Siberian tree line ecotone. Using a space-time approach we are able to exclude the influence of geography and elucidate that lake environmental variables primarily shape the assemblies. We conclude that some diatom genera have greater capabilities to adapt to environmental changes, whereas others will be putatively replaced or lost due to the displacement of the Arctic tundra biome under recent global warming.
Landscapes in high northern latitudes are assumed to be highly sensitive to future global change, but the rates and long-term trajectories of changes are rather uncertain. In the boreal zone, fires are an important factor in climate-vegetation interactions and biogeochemical cycles. Fire regimes are characterized by small, frequent, low-intensity fires within summergreen boreal forests dominated by larch, whereas evergreen boreal forests dominated by spruce and pine burn large areas less frequently but at higher intensities. Here, we explore the potential of the monosaccharide anhydrides (MA) levoglucosan, mannosan and galactosan to serve as proxies of low-intensity biomass burning in glacial-to-interglacial lake sediments from the high northern latitudes. We use sediments from Lake El'gygytgyn (cores PG 1351 and ICDP 5011-1), located in the far north-east of Russia, and study glacial and interglacial samples of the last 430 kyr (marine isotope stages 5e, 6, 7e, 8, 11c and 12) that had different climate and biome configurations. Combined with pollen and non-pollen palynomorph records from the same samples, we assess how far the modern relationships between fire, climate and vegetation persisted during the past, on orbital to centennial timescales. We find that MAs attached to particulates were well-preserved in up to 430 kyr old sediments with higher influxes from low-intensity biomass burning in interglacials compared to glacials. MA influxes significantly increase when summergreen boreal forest spreads closer to the lake, whereas they decrease when tundra-steppe environments and, especially, Sphagnum peatlands spread. This suggests that low-temperature fires are a typical characteristic of Siberian larch forests also on long timescales. The results also suggest that low-intensity fires would be reduced by vegetation shifts towards very dry environments due to reduced biomass availability, as well as by shifts towards peatlands, which limits fuel dryness. In addition, we observed very low MA ratios, which we interpret as high contributions of galactosan and mannosan from biomass sources other than those currently monitored, such as the moss-lichen mats in the understorey of the summergreen boreal forest. Overall, sedimentary MAs can provide a powerful proxy for fire regime reconstructions and extend our knowledge of long-term natural fire-climate-vegetation feedbacks in the high northern latitudes.
Landscapes in high northern latitudes are assumed to be highly sensitive to future global change, but the rates and long-term trajectories of changes are rather uncertain. In the boreal zone, fires are an important factor in climate-vegetation interactions and biogeochemical cycles. Fire regimes are characterized by small, frequent, low-intensity fires within summergreen boreal forests dominated by larch, whereas evergreen boreal forests dominated by spruce and pine burn large areas less frequently but at higher intensities. Here, we explore the potential of the monosaccharide anhydrides (MA) levoglucosan, mannosan and galactosan to serve as proxies of low-intensity biomass burning in glacial-to-interglacial lake sediments from the high northern latitudes. We use sediments from Lake El'gygytgyn (cores PG 1351 and ICDP 5011-1), located in the far north-east of Russia, and study glacial and interglacial samples of the last 430 kyr (marine isotope stages 5e, 6, 7e, 8, 11c and 12) that had different climate and biome configurations. Combined with pollen and non-pollen palynomorph records from the same samples, we assess how far the modern relationships between fire, climate and vegetation persisted during the past, on orbital to centennial timescales. We find that MAs attached to particulates were well-preserved in up to 430 kyr old sediments with higher influxes from low-intensity biomass burning in interglacials compared to glacials. MA influxes significantly increase when summergreen boreal forest spreads closer to the lake, whereas they decrease when tundra-steppe environments and, especially, Sphagnum peatlands spread. This suggests that low-temperature fires are a typical characteristic of Siberian larch forests also on long timescales. The results also suggest that low-intensity fires would be reduced by vegetation shifts towards very dry environments due to reduced biomass availability, as well as by shifts towards peatlands, which limits fuel dryness. In addition, we observed very low MA ratios, which we interpret as high contributions of galactosan and mannosan from biomass sources other than those currently monitored, such as the moss-lichen mats in the understorey of the summergreen boreal forest. Overall, sedimentary MAs can provide a powerful proxy for fire regime reconstructions and extend our knowledge of long-term natural fire-climate-vegetation feedbacks in the high northern latitudes.
Issue Despite their rather similar climatic conditions, eastern Eurasia and northern North America are largely covered by different plant functional types (deciduous or evergreen boreal forest) composed of larch or pine, spruce and fir, respectively. I propose that these deciduous and evergreen boreal forests represent alternative quasi-stable states, triggered by their different northern tree refugia that reflect the different environmental conditions experienced during the Last Glacial. Evidence This view is supported by palaeoecological and environmental evidence. Once established, Asian larch forests are likely to have stabilized through a complex vegetation-fire-permafrost soil-climate feedback system. Conclusion With respect to future forest developments, this implies that Asian larch forests are likely to be governed by long-term trajectories and are therefore largely resistant to natural climate variability on time-scales shorter than millennia. The effects of regional human impact and anthropogenic global warming might, however, cause certain stability thresholds to be crossed, meaning that irreversible transitions occur and resulting in marked consequences for ecosystem services on these human-relevant time-scales.
Pollen-based quantitative land-cover reconstruction for northern Asia covering the last 40 ka cal BP
(2019)
We collected the available relative pollen productivity estimates (PPEs) for 27 major pollen taxa from Eurasia and applied them to estimate plant abundances during the last 40 ka cal BP (calibrated thousand years before present) using pollen counts from 203 fossil pollen records in northern Asia (north of 40 degrees N). These pollen records were organized into 42 site groups and regional mean plant abundances calculated using the REVEALS (Regional Estimates of Vegetation Abundance from Large Sites) model. Time-series clustering, constrained hierarchical clustering, and detrended canonical correspondence analysis were performed to investigate the regional pattern, time, and strength of vegetation changes, respectively. Reconstructed regional plant functional type (PFT) components for each site group are generally consistent with modern vegetation in that vegetation changes within the regions are characterized by minor changes in the abundance of PFTs rather than by an increase in new PFTs, particularly during the Holocene. We argue that pollen-based REVEALS estimates of plant abundances should be a more reliable reflection of the vegetation as pollen may overestimate the turnover, particularly when a high pollen producer invades areas dominated by low pollen producers. Comparisons with vegetation-independent climate records show that climate change is the primary factor driving land-cover changes at broad spatial and temporal scales. Vegetation changes in certain regions or periods, however, could not be explained by direct climate change, e.g. inland Siberia, where a sharp increase in evergreen conifer tree abundance occurred at ca. 7-8 ka cal BP despite an unchanging climate, potentially reflecting their response to complex climate-permafrost-fire-vegetation interactions and thus a possible long-term lagged climate response.
Ongoing and past biome transitions are generally assigned to climate and atmospheric changes (e.g. temperature, precipitation, CO2), but the major regional factors or factor combinations that drive vegetation change often remain unknown. Modelling studies applying ensemble runs can help to partition the effects of the different drivers. Such studies require careful validation with observational data. In this study, fossil pollen records from 741 sites in Europe, 728 sites in North America, and 418 sites in Asia (extracted from terrestrial archives including lake sediments) are used to reconstruct biomes at selected time slices between 40 cal ka BP (calibrated thousand years before present) and today. These results are used to validate Northern Hemisphere biome distributions (>30 degrees N) simulated by the biome model BIOME4 that has been forced with climate data simulated by a General Circulation model. Quantitative comparisons between pollen- and model-based results show a generally good fit at a broad spatial scale. Mismatches occur in central-arid Asia with a broader extent of grassland throughout the last 40 ka (likely due to the over-representation of Artemisia and Chenopodiaceae pollen) and in Europe with over-estimation of tundra at 0 cal ka BP (likely due to human impacts to some extent). Sensitivity analysis reveals that broad-scale biome changes follow the global signal of major postglacial temperature change, although the climatic variables vary in their regional and temporal importance. Temperature is the dominant variable in Europe and other rather maritime areas for biome changes between 21 and 14 ka, while precipitation is highly important in the arid inland regions of Asia and North America. The ecophysiological effect of changes in the atmospheric CO2-concentration has the highest impact during this transition than in other intervals. With respect to modern vegetation in the course of global warming, our findings imply that vegetation change in the Northern Hemisphere may be strongly limited by effective moisture changes, i.e. the combined effect of temperature and precipitation, particularly in inland areas. (C) 2019 Elsevier Ltd. All rights reserved.
Proxy-based reconstructions and modeling of Holocene spatiotemporal precipitation patterns for China and Mongolia have hitherto yielded contradictory results indicating that the basic mechanisms behind the East Asian Summer Monsoon and its interaction with the westerly jet stream remain poorly understood. We present quantitative reconstructions of Holocene precipitation derived from 101 fossil pollen records and analyse them with the help of a minimal empirical model. We show that the westerly jet-stream axis shifted gradually southward and became less tilted since the middle Holocene. This was tracked by the summer monsoon rain band resulting in an early-Holocene precipitation maximum over most of western China, a mid-Holocene maximum in north-central and northeastern China, and a late-Holocene maximum in southeastern China. Our results suggest that a correct simulation of the orientation and position of the westerly jet stream is crucial to the reliable prediction of precipitation patterns in China and Mongolia.
Encroachment of shrubs into the unique pastoral grassland ecosystems of the Tibetan Plateau has significant impact on ecosystem services, especially forage production. We developed a process-based ecohydrological model to identify the relative importance of the main drivers of shrub encroachment for the alpine meadows within the Qinghai province. Specifically, we explored the effects of summer livestock grazing (intensity and type of livestock) together with the effects of climate warming, including interactions between herbaceous and woody vegetation and feedback loops between soil, water and vegetation. Under current climatic conditions and a traditional herd composition, an increasing grazing intensity above a threshold value of 0.32 +/- 0.10 large stock units (LSU) ha(-1) day(-1) changes the vegetation composition from herbaceous towards a woody and bare soil dominated system. Very high grazing intensity (above 0.8 LSU ha(-1) day(-1)) leads to a complete loss of any vegetation. Under warmer conditions, the vegetation showed a higher resilience against livestock farming. This resilience is enhanced when the herd has a higher browser : grazer ratio. A cooler climate has a shrub encroaching effect, whereas warmer conditions increase the cover of the herbaceous vegetation. This effect was primarily due to season length and an accompanied competitive loss of slower growing shrubs, rather than evaporative water loss leading to less soil water in deeper soil layers for deeper rooting shrubs. If climate warming is driving current shrub encroachment, we conclude it is only indirectly so. It would be manifest by an advancing shrubline and could be regarded as a climatic escape of specific shrub species such as Potentilla fruticosa. Under the recent high intensity of grazing, only herding by more browsing animals can potentially prevent both shrub encroachment and the complete loss of herbaceous vegetation.
A strong temperature increase in the Arctic is expected to lead to latitudinal treeline shift. This tundra-taiga turnover would cause a positive vegetation-climate feedback due to albedo decrease. However, reliable estimates of tree migration rates are currently lacking due to the complex processes involved in forest establishment, which depend strongly on seed dispersal. We aim to fill this gap using LAVESI, an individual-based and spatially explicit Larix vegetation simulator. LAVESI was designed to simulate plots within homogeneous forests. Here, we improve the implementation of the seed dispersal function via field-based investigations. We inferred the effective seed dispersal distances of a typical open-forest stand on the southern Taymyr Peninsula (northern central Siberia) from genetic parentage analysis using eight nuclear microsatellite markers. The parentage analysis gives effective seed dispersal distances (median similar to 10 m) close to the seed parents. A comparison between simulated and observed effective seed dispersal distances reveals an overestimation of recruits close to the releasing tree and a shorter dispersal distance generally. We thus adapted our model and used the newly parameterised version to simulate south-to-north transects; a slow-moving treeline front was revealed. The colonisation of the tundra areas was assisted by occasional long-distance seed dispersal events beyond the treeline area. The treeline (similar to 1 tree ha(-1)) advanced by similar to 1.6 m yr(-1), whereas the forest line (similar to 100 trees ha(-1)) advanced by only similar to 0.6 m yr(-1). We conclude that the treeline in northern central Siberia currently lags behind the current strong warming and will continue to lag in the near future.
Larix populations at the tundra-taiga ecotone in northern Siberia are highly under-represented in population genetic studies, possibly due to the remoteness of these regions that can only be accessed at extraordinary expense. The genetic signatures of populations in these boundary regions are therefore largely unknown. We aim to generate organelle reference genomes for the detection of single nucleotide polymorphisms (SNPs) that can be used for paleogenetic studies. We present 19 complete chloroplast genomes and mitochondrial genomic sequences of larches from the southern lowlands of the Taymyr Peninsula (northernmost range of Larix gmelinii (Rupr.) Kuzen.), the lower Omoloy River, and the lower Kolyma River (both in the range of Larix cajanderi Mayr). The genomic data reveal 84 chloroplast SNPs and 213 putatively mitochondrial SNPs. Parsimony-based chloroplast haplotype networks show no spatial structure of individuals from different geographic origins, while the mitochondrial haplotype network shows at least a slight spatial structure with haplotypes from the Omoloy and Kolyma populations being more closely related to each other than to most of the haplotypes from the Taymyr populations. Whole genome alignments with publicly available complete chloroplast genomes of different Larix species show that among official plant barcodes only the rcbL gene contains sufficient polymorphisms, but has to be sequenced completely to distinguish the different provenances. We provide 8 novel mitochondrial SNPs that are putatively diagnostic for the separation of L. gmelinii and L. cajanderi, while 4 chloroplast SNPs have the potential to distinguish the L. gmelinii/ L. cajanderi group from other Larix species. Our organelle references can be used for a targeted primer and probe design allowing the generation of short amplicons. This is particularly important with regard to future investigations of, for example, the biogeographic history of Larix by screening ancient sedimentary DNA of Larix.
Shrub encroachment has far-reaching ecological and economic consequences in many ecosystems worldwide. Yet, compositional changes associated with shrub encroachment are often overlooked despite having important effects on ecosystem functioning. We document the compositional change and potential drivers for a northern Namibian Combretum woodland transitioning into a Terminalia shrubland. We use a multiproxy record (pollen, sedimentary ancient DNA, biomarkers, compound-specific carbon (delta C-13) and deuterium (delta D) isotopes, bulk carbon isotopes (delta(13)Corg), grain size, geochemical properties) from Lake Otjikoto at high taxonomical and temporal resolution. We provide evidence that state changes in semiarid environments may occur on a scale of one century and that transitions between stable states can span around 80 years and are characterized by a unique vegetation composition. We demonstrate that the current grass/woody ratio is exceptional for the last 170 years, as supported by n-alkane distributions and the delta C-13 and delta(13)Corg records. Comparing vegetation records to environmental proxy data and census data, we infer a complex network of global and local drivers of vegetation change. While our delta D record suggests physiological adaptations of woody species to higher atmospheric pCO(2) concentration and drought, our vegetation records reflect the impact of broad-scale logging for the mining industry, and the macrocharcoal record suggests a decrease in fire activity associated with the intensification of farming. Impact of selective grazing is reflected by changes in abundance and taxonomical composition of grasses and by an increase of nonpalatable and trampling-resistant taxa. In addition, grain-size and spore records suggest changes in the erodibility of soils because of reduced grass cover. Synthesis. We conclude that transitions to an encroached savanna state are supported by gradual environmental changes induced by management strategies, which affected the resilience of savanna ecosystems. In addition, feedback mechanisms that reflect the interplay between management legacies and climate change maintain the encroached state.
Forest structure is a crucial component in the assessment of whether a forest is likely to act as a carbon sink under changing climate. Detailed 3D structural information about the tundra–taiga ecotone of Siberia is mostly missing and still underrepresented in current research due to the remoteness and restricted accessibility. Field based, high-resolution remote sensing can provide important knowledge for the understanding of vegetation properties and dynamics. In this study, we test the applicability of consumer-grade Unmanned Aerial Vehicles (UAVs) for rapid calculation of stand metrics in treeline forests. We reconstructed high-resolution photogrammetric point clouds and derived canopy height models for 10 study sites from NE Chukotka and SW Yakutia. Subsequently, we detected individual tree tops using a variable-window size local maximum filter and applied a marker-controlled watershed segmentation for the delineation of tree crowns. With this, we successfully detected 67.1% of the validation individuals. Simple linear regressions of observed and detected metrics show a better correlation (R2) and lower relative root mean square percentage error (RMSE%) for tree heights (mean R2 = 0.77, mean RMSE% = 18.46%) than for crown diameters (mean R2 = 0.46, mean RMSE% = 24.9%). The comparison between detected and observed tree height distributions revealed that our tree detection method was unable to representatively identify trees <2 m. Our results show that plot sizes for vegetation surveys in the tundra–taiga ecotone should be adapted to the forest structure and have a radius of >15–20 m to capture homogeneous and representative forest stands. Additionally, we identify sources of omission and commission errors and give recommendations for their mitigation. In summary, the efficiency of the used method depends on the complexity of the forest’s stand structure.
Forest structure is a crucial component in the assessment of whether a forest is likely to act as a carbon sink under changing climate. Detailed 3D structural information about the tundra–taiga ecotone of Siberia is mostly missing and still underrepresented in current research due to the remoteness and restricted accessibility. Field based, high-resolution remote sensing can provide important knowledge for the understanding of vegetation properties and dynamics. In this study, we test the applicability of consumer-grade Unmanned Aerial Vehicles (UAVs) for rapid calculation of stand metrics in treeline forests. We reconstructed high-resolution photogrammetric point clouds and derived canopy height models for 10 study sites from NE Chukotka and SW Yakutia. Subsequently, we detected individual tree tops using a variable-window size local maximum filter and applied a marker-controlled watershed segmentation for the delineation of tree crowns. With this, we successfully detected 67.1% of the validation individuals. Simple linear regressions of observed and detected metrics show a better correlation (R2) and lower relative root mean square percentage error (RMSE%) for tree heights (mean R2 = 0.77, mean RMSE% = 18.46%) than for crown diameters (mean R2 = 0.46, mean RMSE% = 24.9%). The comparison between detected and observed tree height distributions revealed that our tree detection method was unable to representatively identify trees <2 m. Our results show that plot sizes for vegetation surveys in the tundra–taiga ecotone should be adapted to the forest structure and have a radius of >15–20 m to capture homogeneous and representative forest stands. Additionally, we identify sources of omission and commission errors and give recommendations for their mitigation. In summary, the efficiency of the used method depends on the complexity of the forest’s stand structure.
It is of major interest to estimate the feedback of arctic ecosystems to the global warming we expect in upcoming decades. The speed of this response is driven by the potential of species to migrate, tracking their climate optimum. For this, sessile plants have to produce and disperse seeds to newly available habitats, and pollination of ovules is needed for the seeds to be viable. These two processes are also the vectors that pass genetic information through a population. A restricted exchange among subpopulations might lead to a maladapted population due to diversity losses. Hence, a realistic implementation of these dispersal processes into a simulation model would allow an assessment of the importance of diversity for the migration of plant species in various environments worldwide. To date, dynamic global vegetation models have been optimized for a global application and overestimate the migration of biome shifts in currently warming temperatures. We hypothesize that this is caused by neglecting important fine-scale processes, which are necessary to estimate realistic vegetation trajectories. Recently, we built and parameterized a simulation model LAVESI for larches that dominate the latitudinal treelines in the northernmost areas of Siberia. In this study, we updated the vegetation model by including seed and pollen dispersal driven by wind speed and direction. The seed dispersal is modelled as a ballistic flight, and for the pollination of ovules of seeds produced, we implemented a wind-determined and distance-dependent probability distribution function using a von Mises distribution to select the pollen donor. A local sensitivity analysis of both processes supported the robustness of the model's results to the parameterization, although it highlighted the importance of recruitment and seed dispersal traits for migration rates. This individual-based and spatially explicit implementation of both dispersal processes makes it easily feasible to inherit plant traits and genetic information to assess the impact of migration processes on the genetics. Finally, we suggest how the final model can be applied to substantially help in unveiling the important drivers of migration dynamics and, with this, guide the improvement of recent global vegetation models.
Recent global warming is pronounced in high-latitude regions (e.g. northern Asia), and will cause the vegetation to change. Future vegetation trends (e.g. the "arctic greening") will feed back into atmospheric circulation and the global climate system. Understanding the nature and causes of past vegetation changes is important for predicting the composition and distribution of future vegetation communities. Fossil pollen records from 468 sites in northern and eastern Asia were biomised at selected times between 40 cal ka bp and today. Biomes were also simulated using a climate-driven biome model and results from the two approaches compared in order to help understand the mechanisms behind the observed vegetation changes. The consistent biome results inferred by both approaches reveal that long-term and broad-scale vegetation patterns reflect global- to hemispheric-scale climate changes. Forest biomes increase around the beginning of the late deglaciation, become more widespread during the early and middle Holocene, and decrease in the late Holocene in fringe areas of the Asian Summer Monsoon. At the southern and southwestern margins of the taiga, forest increases in the early Holocene and shows notable species succession, which may have been caused by winter warming at ca. 7 cal ka bp. At the northeastern taiga margin (central Yakutia and northeastern Siberia), shrub expansion during the last deglaciation appears to prevent the permafrost from thawing and hinders the northward expansion of evergreen needle-leaved species until ca. 7 cal ka bp. The vegetation-climate disequilibrium during the early Holocene in the taiga-tundra transition zone suggests that projected climate warming will not cause a northward expansion of evergreen needle-leaved species.
It is of major interest to estimate the feedback of arctic ecosystems to the global warming we expect in upcoming decades. The speed of this response is driven by the potential of species to migrate, tracking their climate optimum. For this, sessile plants have to produce and disperse seeds to newly available habitats, and pollination of ovules is needed for the seeds to be viable. These two processes are also the vectors that pass genetic information through a population. A restricted exchange among subpopulations might lead to a maladapted population due to diversity losses. Hence, a realistic implementation of these dispersal processes into a simulation model would allow an assessment of the importance of diversity for the migration of plant species in various environments worldwide. To date, dynamic global vegetation models have been optimized for a global application and overestimate the migration of biome shifts in currently warming temperatures. We hypothesize that this is caused by neglecting important fine-scale processes, which are necessary to estimate realistic vegetation trajectories. Recently, we built and parameterized a simulation model LAVESI for larches that dominate the latitudinal treelines in the northernmost areas of Siberia. In this study, we updated the vegetation model by including seed and pollen dispersal driven by wind speed and direction. The seed dispersal is modelled as a ballistic flight, and for the pollination of ovules of seeds produced, we implemented a wind-determined and distance-dependent probability distribution function using a von Mises distribution to select the pollen donor. A local sensitivity analysis of both processes supported the robustness of the model's results to the parameterization, although it highlighted the importance of recruitment and seed dispersal traits for migration rates. This individual-based and spatially explicit implementation of both dispersal processes makes it easily feasible to inherit plant traits and genetic information to assess the impact of migration processes on the genetics. Finally, we suggest how the final model can be applied to substantially help in unveiling the important drivers of migration dynamics and, with this, guide the improvement of recent global vegetation models.
Understanding vegetation-modern pollen relationships is essential to provide confidence in fossil pollen reconstructions of long-term vegetation changes in savanna ecosystems. In this paper we compare the taxonomical composition and the diversity (Hill NO, N1, N2) of vegetation and modern pollen along precipitation and local grazing-intensity gradients in Namibian savannas. Modern pollen was extracted from surface soil samples collected from 5 x 5 m plots distributed along four 500 m gradients. Vegetation was surveyed in each plot. The results show a high correspondence between vegetation and pollen data in terms of composition. Precipitation and grazing explain a significant although low proportion of compositional change in the vegetation and pollen spectra. We identified pollen taxa as indicators of grazing pressure such as Limeum, Alternanthera, and particularly Tribulus. Correspondence between vegetation and pollen data in terms of taxa richness (NO) is limited, probably because of the influence of landscape heterogeneity and openness, as well as low pollen concentrations. In contrast, the effective numbers of common and dominant taxa (N1, N2) are consistent among the different datasets. We conclude that in spite of limitations, modern pollen assemblages can reflect changes in vegetation composition, richness and diversity patterns along precipitation and grazing gradients in savanna environments. (C) 2018 Elsevier B.V. All rights reserved.
The climate conditions during Marine Isotope Stage (MIS) 3 were similar to present-day conditions, but whether humidity then exceeded present levels is debated, and the driving mechanisms of palaeoclimate evolution since MIS 3 remain unclear. Here, we use pollen data from Wulagai Lake, Inner Mongolia, to reconstruct vegetation and climate changes since the middle MIS 3. The steppe biome is reconstructed as the first dominant biome and the desert biome as the second, and the results show that the vegetation was steppe over the last 43,800 years. Poaceae, Artemisia, Caryophyllaceae and Humulus were abundant from middle to late MIS 3, indicating humid climate conditions. As drought-tolerant species such as Hippophae, Nitraria and Chenopodiaceae spread during MIS 2, the climate became arid. The Holocene is characterized by the dominance of steppe with mixed coniferous-broadleaved forests in the Greater Hinggan Range, and the desert biome retains high affinity scores, indicating that the climate was semi-arid. The climate from middle to late MIS 3 was wetter than in the Holocene; this shift was related to changes in the Northern Hemisphere's solar insolation and ice volume. The humid conditions during MIS 3 were attributed to strong ice–albedo feedback, which led to evaporation that was less than the precipitation. The enhanced evaporation caused by increased solar insolation and decreased ice volume might have exceeded the precipitation during the Holocene and resulted in low effective humidity in the Wulagai Lake basin.
We present a pollen record for last 28 cal kyr BP from the eastern basin of Lake Karakul, the largest lake in Tajikistan, located in the eastern Pamir Mountains at 3915 m asl, a geographically complex region. The pollen record is dominated by Artemisia and Chenopodiaceae, while other taxa, apart from Poaceae, are present in low quantities and rarely exceed 5% in total. Arboreal pollen occur predominantly from similar to 28 to similar to 13 cal kyr BP, but as likely no trees occurred in the high mountain regions of the eastern Pamir during this time due to the high altitude and cold climate, arboreal taxa are attributed to long distance transport, probably by the Westerlies, the dominant atmospheric circulation. Tree pollen influx decreases strongly after similar to 13 cal kyr BP, allowing the pollen spectra to be interpreted as a regional vegetation signal. We infer that from 27.6 to 19.4 cal kyr BP the eastern Pamir was dominated by dry mountain steppe with low vegetation cover, while from 19.0 to 13.6 cal kyr BP Artemisia values increase and Chenopodiaceae, most herb taxa, and inferred far distant input from arboreal taxa decrease. Between 12.9 and 6.7 cal kyr BP open steppe vegetation dominated with maximum values in Ephedra, and while steppe taxa still dominated the spectra from 5.4 to 1 cal kyr BP, meadow taxa start to increase. During the last millennium, alpine steppe and alpine meadows expanded and a weak human influence can be ascertained from the increase of Asteraceae and the occurrence of Plantago pollen in the spectra.
In this study, we analyze interactions in lake and lake catchment systems of a continuous permafrost area. We assessed colored dissolved organic matter (CDOM) absorption at 440 nm (a(440)(CDOM)) and absorption slope (S300-500) in lakes using field sampling and optical remote sensing data for an area of 350 km(2) in Central Yamal, Siberia. Applying a CDOM algorithm (ratio of green and red band reflectance) for two high spatial resolution multispectral GeoEye-1 and Worldview-2 satellite images, we were able to extrapolate the a()(CDOM) data from 18 lakes sampled in the field to 356 lakes in the study area (model R-2 = 0.79). Values of a(440)(CDOM) in 356 lakes varied from 0.48 to 8.35 m(-1) with a median of 1.43 m(-1). This a()(CDOM) dataset was used to relate lake CDOM to 17 lake and lake catchment parameters derived from optical and radar remote sensing data and from digital elevation model analysis in order to establish the parameters controlling CDOM in lakes on the Yamal Peninsula. Regression tree model and boosted regression tree analysis showed that the activity of cryogenic processes (thermocirques) in the lake shores and lake water level were the two most important controls, explaining 48.4% and 28.4% of lake CDOM, respectively (R-2 = 0.61). Activation of thermocirques led to a large input of terrestrial organic matter and sediments from catchments and thawed permafrost to lakes (n = 15, mean a(440)(CDOM) = 5.3 m(-1)). Large lakes on the floodplain with a connection to Mordy-Yakha River received more CDOM (n = 7, mean a(440)(CDOM) = 3.8 m(-1)) compared to lakes located on higher terraces.
In this study, we analyze interactions in lake and lake catchment systems of a continuous permafrost area. We assessed colored dissolved organic matter (CDOM) absorption at 440 nm (a(440)(CDOM)) and absorption slope (S300-500) in lakes using field sampling and optical remote sensing data for an area of 350 km(2) in Central Yamal, Siberia. Applying a CDOM algorithm (ratio of green and red band reflectance) for two high spatial resolution multispectral GeoEye-1 and Worldview-2 satellite images, we were able to extrapolate the a()(CDOM) data from 18 lakes sampled in the field to 356 lakes in the study area (model R-2 = 0.79). Values of a(440)(CDOM) in 356 lakes varied from 0.48 to 8.35 m(-1) with a median of 1.43 m(-1). This a()(CDOM) dataset was used to relate lake CDOM to 17 lake and lake catchment parameters derived from optical and radar remote sensing data and from digital elevation model analysis in order to establish the parameters controlling CDOM in lakes on the Yamal Peninsula. Regression tree model and boosted regression tree analysis showed that the activity of cryogenic processes (thermocirques) in the lake shores and lake water level were the two most important controls, explaining 48.4% and 28.4% of lake CDOM, respectively (R-2 = 0.61). Activation of thermocirques led to a large input of terrestrial organic matter and sediments from catchments and thawed permafrost to lakes (n = 15, mean a(440)(CDOM) = 5.3 m(-1)). Large lakes on the floodplain with a connection to Mordy-Yakha River received more CDOM (n = 7, mean a(440)(CDOM) = 3.8 m(-1)) compared to lakes located on higher terraces.
Changes in species' distributions are classically projected based on their climate envelopes. For Siberian forests, which have a tremendous significance for vegetation-climate feedbacks, this implies future shifts of each of the forest-forming larch (Larix) species to the north-east. However, in addition to abiotic factors, reliable projections must assess the role of historical biogeography and biotic interactions. Here, we use sedimentary ancient DNA and individual-based modelling to investigate the distribution of larch species and mitochondrial haplotypes through space and time across the treeline ecotone on the southern Taymyr peninsula, which at the same time presents a boundary area of two larch species. We find spatial and temporal patterns, which suggest that forest density is the most influential driver determining the precise distribution of species and mitochondrial haplotypes. This suggests a strong influence of competition on the species' range shifts. These findings imply possible climate change outcomes that are directly opposed to projections based purely on climate envelopes. Investigations of such fine-scale processes of biodiversity change through time are possible using paleoenvironmental DNA, which is available much more readily than visible fossils and can provide information at a level of resolution that is not reached in classical palaeoecology.
Changes in species’ distributions are classically projected based on their climate envelopes. For Siberian forests, which have a tremendous significance for vegetation-climate feedbacks, this implies future shifts of each of the forest-forming larch (Larix) species to the north-east. However, in addition to abiotic factors, reliable projections must assess the role of historical biogeography and biotic interactions. Here, we use sedimentary ancient DNA and individual-based modelling to investigate the distribution of larch species and mitochondrial haplotypes through space and time across the treeline ecotone on the southern Taymyr peninsula, which at the same time presents a boundary area of two larch species. We find spatial and temporal patterns, which suggest that forest density is the most influential driver determining the precise distribution of species and mitochondrial haplotypes. This suggests a strong influence of competition on the species’ range shifts. These findings imply possible climate change outcomes that are directly opposed to projections based purely on climate envelopes. Investigations of such fine-scale processes of biodiversity change through time are possible using paleoenvironmental DNA, which is available much more readily than visible fossils and can provide information at a level of resolution that is not reached in classical palaeoecology.
Arctic treelines are facing a strong temperature increase as a result of recent global warming, causing possible changes in forest extent, which will alter vegetation-climate feedbacks. However, the mode and strength of the response is rather unclear, as potential changes are happening in areas that are very remote and difficult to access, and empirical data are still largely lacking. Here, we assessed the current population structure and genetic differentiation of Larix Mill. tree stands within the northernmost latitudinal treeline reaching ~ 72° N in the southern lowlands of the Taymyr Peninsula (~ 100° E). We sampled 743 individuals belonging to different height classes (seedlings, saplings, trees) at 11 locations along a gradient from ‘single tree’ tundra over ‘forest line’ to ‘dense forest’ stands and conducted investigations applying eight highly polymorphic nuclear microsatellites. Results suggest a high diversity within sub-populations (HE = 0.826–0.893), coupled, however, with heterozygote deficits in all sub-populations, but pronounced in ‘forest line’ stands. Overall, genetic differentiation of sub-populations is low (FST = 0.005), indicating a region-wide high gene flow, although ‘forest line’ stands harbour few rare and private alleles, likely indicating greater local reproduction. ‘Single tree’ stands, located beyond the northern forest line, are currently not involved in treeline expansion, but show signs of a long-term refuge, namely asexual reproduction and change of growth-form from erect to creeping growth, possibly having persisted for thousands of years. The lack of differentiation between the sub-populations points to a sufficiently high dispersal potential, and thus a rapid northward migration of the Siberian arctic treeline under recent global warming seems potentially unconstrained, but observations show it to be unexpectedly slow.