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Siberian arctic vegetation and lake water communities, known for their temperature dependence, are expected to be particularly impacted by recent climate change and high warming rates. However, decadal information on the nature and strength of recent vegetation change and its time lag to climate signals are rare. In this study, we present a Pb-210/Cs-137 dated pollen and Pediastrum species record from a unnamed lake in the south of the Taymyr peninsula covering the period from AD 1706 to 2011. Thirty-nine palynomorphs and 10 morphotypes of Pediastrum species were studied to assess changes in vegetation and lake conditions as probable responses to climate change. We compared the pollen record with Pediastrum species, which we consider to be important proxies of climate changes. Three pollen assemblage zones characterised by Betula nana, Alnus viridis and Larix gmelinii (1706-1808); herbs such as Cyperaceae, Artemisia or Senecio (1808-1879), and higher abundance of Larix pollen (1955-2011) are visible. Also, three Pediastrum assemblage zones show changes of aquatic conditions: higher abundances of Pediastrum boryanum var. brevicorne (1706-1802); medium abundances of P. kawraiskyi and P. integrum (1802-1840 and 1920-1980), indicating cooler conditions while less eutrophic conditions are indicated by P. boryanum, and a mainly balanced composition with only small changes of cold- and warm-adapted Pediastrum species (1965-2011). In general, compositional Pediastrum species turnover is slightly higher than that indicated by pollen data (0.54 vs 0.34 SD), but both are only minor for this treeline location. In conclusion, the relevance of differentiation of Pediastrum species is promising and can give further insights into the relationship between lakes and their surrounding vegetation transferred onto climatic conditions.
Reliable information on past and present vegetation is important to project future changes, especially for rapidly transitioning areas such as the boreal treeline. To study past vegetation, pollen analysis is common, while current vegetation is usually assessed by field surveys. Application of detailed sedimentary DNA (sedDNA) records has the potential to enhance our understanding of vegetation changes, but studies systematically investigating the power of this proxy are rare to date. This study compares sedDNA metabarcoding and pollen records from surface sediments of 31 lakes along a north-south gradient of increasing forest cover in northern Siberia (Taymyr peninsula) with data from field surveys in the surroundings of the lakes. sedDNA metabarcoding recorded 114 plant taxa, about half of them to species level, while pollen analyses identified 43 taxa, both exceeding the 31 taxa found by vegetation field surveys. Increasing Larix percentages from north to south were consistently recorded by all three methods and principal component analyses based on percentage data of vegetation surveys and DNA sequences separated tundra from forested sites. Comparisons of the ordinations using procrustes and protest analyses show a significant fit among all compared pairs of records. Despite similarities of sedDNA and pollen records, certain idiosyncrasies, such as high percentages of Alnus and Betula in all pollen and high percentages of Salix in all sedDNA spectra, are observable. Our results from the tundra to single-tree tundra transition zone show that sedDNA analyses perform better than pollen in recording site-specific richness (i.e., presence/absence of taxa in the vicinity of the lake) and perform as well as pollen in tracing vegetation composition.
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.
Die vorliegende Arbeit wurde im Rahmen des multidisziplinären Deutsch-Russischen Verbundprojektes "Laptev See 2000" erstellt. Die dargestellten bodenkundlichen und mikro-biologischen Untersuchungen verfolgten das Ziel die mikrobielle Lebensgemeinschaft eines Permafrostbodens im sibirischen Lena Delta zu charakterisieren, wobei den methanogenen Archaea besondere Beachtung zukam. Die Probennahme wurde im August 2001 im zentralen Lenadelta, auf der Insel Samoylov durchgeführt. Das Delta liegt im Bereich des kontinuierlichen Permafrostes, was bedeutet, dass nur eine flache saisonale Auftauschicht während der Sommermonate auftaut. Das untersuchte Bodenprofil lag im Zentrum eines für die Landschaft repräsentativen Low Center Polygons. Zum Zeitpunkt der Beprobung betrug die Auftautiefe des untersuchten Bodens 45 cm.. Der Wasserstand lag zum Untersuchungszeitpunkt 18 cm unter der Geländeoberfläche, so dass alle tiefer liegenden Horizonte durch anaerobe Verhältnisse charakterisiert waren. Die Untersuchung der bodenkundlichen Parameter ergab unter anderem eine mit zunehmender Tiefe abnehmende Konzentration von Kohlenstoff und Stickstoff, sowie eine Abnahme von Temperatur und Wurzeldichte. Um die Auswirkungen der sich mit der Tiefe verändernden Bodeneigenschaften auf die Mikroorganismen zu ermitteln, wurden die Mikroorganismenpopulationen der verschiedenen Bodentiefen mit Hilfe der Fluoreszenz in situ Hybridisierung hinsichtlich ihrer Anzahl, Aktivität und Zusammensetzung beschrieben. Für die Charakterisierung des physiologischen Profils dieser Gemeinschaften, bezüglich der von ihr umsetzbaren Kohlenstoffverbindungen, wurden BIOLOG Mikrotiterplatten unter den in situ Bedingungen angepassten Bedingungen eingesetzt. Die sich im Profil verändernden Bodenparameter, vor allem die abnehmende Substratversorgung, die geringe Temperatur und die anaeroben Verhältnisse in den unteren Bodenschichten führten zu einer Veränderung der Mikroorganismenpopulation im Bodenprofil. So nahm von oben nach unten die Gesamtanzahl der ermittelten Mikroorganismen von 23,0 × 108 auf 1,2 × 108 Zellen g-1 ab. Gleichzeitig sank der Anteil der aktiven Zellen von 59% auf 33%. Das bedeutet, dass im Bereich von 0-5 cm 35mal mehr aktive Zellen g-1 als im Bereich von 40-45 cm gefunden wurden. Durch den Einsatz spezieller rRNS-Sonden konn-te zusätzlich eine Abnahme der Diversität mit zunehmender Bodentiefe nachgewiesen werden. Die geringere Aktivität der Population in den unteren Horizonten sowie die Unterschiede in der Zusammensetzung wirkten sich auf den Abbau der organischen Substanz aus. So wur-den die mit Hilfe der BIOLOG Mikrotiterplatten angebotenen Substanzen in größerer Tiefe langsamer und unvollständiger abgebaut. Insbesondere in den oberen 5 cm konnten einige der angebotenen Polymere und Kohlehydrate deutlich besser als im restlichen Profil umge-setzt werden. Das außerdem unter anaeroben Versuchsbedingungen diese Substrate deutlich schlechter umgesetzt wurden, kann so interpretiert werden, dass die konstant anaeroben Bedingungen in den unteren Horizonten ein Auftreten der Arten, die diese Substrate umset-zen, erschweren. Die in den oberen, aeroben Bodenabschnitten wesentlich höheren Zellzahlen und Aktivitäten und die dadurch schnellere C-Umsetzung führen auch zu einer besseren Substratversorgung der methanogenen Archaea in den makroskopisch aeroben Horizonten. Die erhöhte Substratverfügbarkeit erklärt die Tatsache, dass im Bereich von 0-5 cm die meisten methanogenen Archaea gefunden wurden, obwohl sich dieser Bereich zum Zeitpunkt der Probennahme oberhalb des wassergesättigten Bodenbereichs befand. Trotz der aeroben Bedingungen in, liegt im Bereich von 5 10 cm die für die methanogenen Archaea am besten geeignete Kombination aus Substratangebot und anaeroben Nischen vor. Hinzu kommt, dass in diesen Tiefen die Sommertemperaturen etwas höher liegen als in den tieferen Horizonten, was wiederum die Aktivität positiv beeinflusst. Bei zusammenfassender Betrachtung der Untersuchungsergebnisse von Anzahl, Aktivität, Zusammensetzung und Leistung der gesamten, aber im besonderen auch der methanogenen Mikroorganismenpopulation wird deutlich, dass in dem untersuchten Bodenprofil unter ökologischen Gesichtspunkten die oberen 15-20 cm den für den C-Umsatz relevantesten Bereich darstellen. Das Zusammenspiel wichtiger Bodenparameter wie Bodentemperatur, Wasserstand, Nährstoffversorgung und Durchwurzelung führt dazu, dass in dem untersuchten Tundraboden in den oberen 15-20 cm eine wesentlich größere und diversere Anzahl an Mikroorganismen existiert, die für einen schnelleren und umfassenderen Kohlenstoffumsatz in diesem Bereich des active layers sorgt.
Woody plants are expanding into the Arctic in response to the warming climate. The impact on arctic plant communities is not well understood due to the limited knowledge about plant assembly rules.
Records of past plant diversity over long time series are rare. Here, we applied sedimentary ancient DNA metabarcoding targeting the P6 loop of the chloroplast trnL gene to a sediment record from Lake Ilirney (central Chukotka, Far Eastern Russia) covering the last 28 thousand years.
Our results show that forb-rich steppe-tundra and dwarf-shrub tundra dominated during the cold climate before 14 ka, while deciduous erect-shrub tundra was abundant during the warm period since 14 ka. Larix invasion during the late Holocene substantially lagged behind the likely warmest period between 10 and 6 ka, where the vegetation biomass could be highest.
We reveal highest richness during 28-23 ka and a second richness peak during 13-9 ka, with both periods being accompanied by low relative abundance of shrubs. During the cold period before 14 ka, rich plant assemblages were phylogenetically clustered, suggesting low genetic divergence in the assemblages despite the great number of species. This probably originates from environmental filtering along with niche differentiation due to limited resources under harsh environmental conditions. In contrast, during the warmer period after 14 ka, rich plant assemblages were phylogenetically overdispersed.
This results from a high number of species which were found to harbor high genetic divergence, likely originating from an erratic recruitment process in the course of warming. Some of our evidence may be of relevance for inferring future arctic plant assembly rules and diversity changes. By analogy to the past, we expect a lagged response of tree invasion. Plant richness might overshoot in the short term; in the long-term, however, the ongoing expansion of deciduous shrubs will eventually result in a phylogenetically more diverse community.
This thesis investigates how the permafrost microbiota responds to global warming. In detail, the constraints behind methane production in thawing permafrost were linked to methanogenic activity, abundance and composition. Furthermore, this thesis offers new insights into microbial adaptions to the changing environmental conditions during global warming. This was assesed by investigating the potential ecological relevant functions encoded by plasmid DNA within the permafrost microbiota. Permafrost of both interglacial and glacial origin spanning the Holocene to the late Pleistocene, including Eemian, were studied during long-term thaw incubations. Furthermore, several permafrost cores of different stratigraphy, soil type and vegetation cover were used to target the main constraints behind methane production during short-term thaw simulations. Short- and long-term incubations simulating thaw with and without the addition of substrate were combined with activity measurements, amplicon and metagenomic sequencing of permanently frozen and seasonally thawed active layer. Combined, it allowed to address the following questions. i) What constraints methane production when permafrost thaws and how is this linked to methanogenic activity, abundance and composition? ii) How does the methanogenic community composition change during long-term thawing conditions? iii) Which potential ecological relevant functions are encoded by plasmid DNA in active layer soils?
The major outcomes of this thesis are as follows. i) Methane production from permafrost after long-term thaw simulation was found to be constrained mainly by the abundance of methanogens and the archaeal community composition. Deposits formed during periods of warmer temperatures and increased precipitation, (here represented by deposits from the Late Pleistocene of both interstadial and interglacial periods) were found to respond strongest to thawing conditions and to contain an archaeal community dominated by methanogenic archaea (40% and 100% of all detected archaea). Methanogenic population size and carbon density were identified as main predictors for potential methane production in thawing permafrost in short-term incubations when substrate was sufficiently available.
ii) Besides determining the methanogenic activity after long-term thaw, the paleoenvironmental conditions were also found to influence the response of the methanogenic community composition. Substantial shifts within methanogenic community structure and a drop in diversity were observed in deposits formed during warmer periods, but not in deposits from stadials, when colder and drier conditions occurred. Overall, a shift towards a dominance of hydrogenotrophic methanogens was observed in all samples, except for the oldest interglacial deposits from the Eemian, which displayed a potential dominance of acetoclastic methanogens. The Eemian, which is discussed to serve as an analogue to current climate conditions, contained highly active methanogenic communities. However, all potential limitation of methane production after permafrost thaw, it means methanogenic community structure, methanogenic population size, and substrate pool might be overcome after permafrost had thawed on the long-term. iii) Enrichments with soil from the seasonally thawed active layer revealed that its plasmid DNA (‘metaplasmidome’) carries stress-response genes. In particular it encoded antibiotic resistance genes, heavy metal resistance genes, cold shock proteins and genes encoding UV-protection. Those are functions that are directly involved in the adaptation of microbial communities to stresses in polar environments. It was further found that metaplasmidomes from the Siberian active layer originate mainly from Gammaproteobacteria. By applying enrichment cultures followed by plasmid DNA extraction it was possible to obtain a higher average contigs length and significantly higher recovery of plasmid sequences than from extracting plasmid sequences from metagenomes. The approach of analyzing ‘metaplasmidomes’ established in this thesis is therefore suitable for studying the ecological role of plasmids in polar environments in general.
This thesis emphasizes that including microbial community dynamics have the potential to improve permafrost-carbon projections. Microbially mediated methane release from permafrost environments may significantly impact future climate change. This thesis identified drivers of methanogenic composition, abundance and activity in thawing permafrost landscapes. Finally, this thesis underlines the importance to study how the current warming Arctic affects microbial communities in order to gain more insight into microbial response and adaptation strategies.
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.
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.
About 24 % of the land surface in the northern hemisphere are underlayed by permafrost in various states. Permafrost aggradation occurs under special environmental conditions with overall low annual precipitation rates and very low mean annual temperatures. Because the general permafrost occurrence is mainly driven by large-scale climatic conditions, the distribution of permafrost deposits can be considered as an important climate indicator. The region with the most extensive continuous permafrost is Siberia. In northeast Siberia, the ice- and organic-rich permafrost deposits of the Ice Complex are widely distributed. These deposits consist mostly of silty to fine-grained sandy sediments that were accumulated during the Late Pleistocene in an extensive plain on the then subaerial Laptev Sea shelf. One important precondition for the Ice Complex sedimentation was, that the Laptev Sea shelf was not glaciated during the Late Pleistocene, resulting in a mostly continuous accumulation of permafrost sediments for at least this period. This shelf landscape became inundated and eroded in large parts by the Holocene marine transgression after the Last Glacial Maximum. Remnants of this landscape are preserved only in the present day coastal areas. Because the Ice Complex deposits contain a wide variety of palaeo-environmental proxies, it is an excellent palaeo-climate archive for the Late Quaternary in the region. Furthermore, the ice-rich Ice Complex deposits are sensible to climatic change, i.e. climate warming. Because of the large-scale climatic changes at the transition from the Pleistocene to the Holocene, the Ice Complex was subject to extensive thermokarst processes since the Early Holocene. Permafrost deposits are not only an environmental indicator, but also an important climate factor. Tundra wetlands, which have developed in environments with aggrading permafrost, are considered a net sink for carbon, as organic matter is stored in peat or is syn-sedimentary frozen with permafrost aggradation. Contrary, the Holocene thermokarst development resulted in permafrost degradation and thus the release of formerly stored organic carbon. Modern tundra wetlands are also considered an important source for the climate-driving gas methane, originating mainly from microbial activity in the seasonal active layer. Most scenarios for future global climate development predict a strong warming trend especially in the Arctic. Consequently, for the understanding of how permafrost deposits will react and contribute to such scenarios, it is necessary to investigate and evaluate ice-rich permafrost deposits like the widespread Ice Complex as climate indicator and climate factor during the Late Quaternary. Such investigations are a pre-condition for the precise modelling of future developments in permafrost distribution and the influence of permafrost degradation on global climate. The focus of this work, which was conducted within the frame of the multi-disciplinary joint German-Russian research projects "Laptev Sea 2000" (1998-2002) and "Dynamics of Permafrost" (2003-2005), was twofold. First, the possibilities of using remote sensing and terrain modelling techniques for the observation of periglacial landscapes in Northeast Siberia in their present state was evaluated and applied to key sites in the Laptev Sea coastal lowlands. The key sites were situated in the eastern Laptev Sea (Bykovsky Peninsula and Khorogor Valley) and the western Laptev Sea (Cape Mamontovy Klyk region). For this task, techniques using CORONA satellite imagery, Landsat-7 satellite imagery, and digital elevation models were developed for the mapping of periglacial structures, which are especially indicative of permafrost degradation. The major goals were to quantify the extent of permafrost degradation structures and their distribution in the investigated key areas, and to establish techniques, which can be used also for the investigation of other regions with thermokarst occurrence. Geographical information systems were employed for the mapping, the spatial analysis, and the enhancement of classification results by rule-based stratification. The results from the key sites show, that thermokarst, and related processes and structures, completely re-shaped the former accumulation plain to a strongly degraded landscape, which is characterised by extensive deep depressions and erosional remnants of the Late Pleistocene surface. As a results of this rapid process, which in large parts happened within a short period during the Early Holocene, the hydrological and sedimentological regime was completely changed on a large scale. These events resulted also in a release of large amounts of organic carbon. Thermokarst is now the major component in the modern periglacial landscapes in terms of spatial extent, but also in its influence on hydrology, sedimentation and the development of vegetation assemblages. Second, the possibilities of using remote sensing and terrain modelling as a supplementary tool for palaeo-environmental reconstructions in the investigated regions were explored. For this task additionally a comprehensive cryolithological field database was developed for the Bykovsky Peninsula and the Khorogor Valley, which contains previously published data from boreholes, outcrops sections, subsurface samples, and subsurface samples, as well as additional own field data. The period covered by this database is mainly the Late Pleistocene and the Holocene, but also the basal deposits of the sedimentary sequence, interpreted as Pliocene to Early Pleistocene, are contained. Remote sensing was applied for the observation of periglacial strucures, which then were successfully related to distinct landscape development stages or time intervals in the investigation area. Terrain modelling was used for providing a general context of the landscape development. Finally, a scheme was developed describing mainly the Late Quaternary landscape evolution in this area. A major finding was the possibility of connecting periglacial surface structures to distinct landscape development stages, and thus use them as additional palaeo-environmental indicator together with other proxies for area-related palaeo-environmental reconstructions. In the landscape evolution scheme, i.e. of the genesis of the Late Pleistocene Ice Complex and the Holocene thermokarst development, some new aspects are presented in terms of sediment source and general sedimentation conditions. This findings apply also for other sites in the Laptev Sea region.
Thermal permafrost degradation and coastal erosion in the Arctic remobilize substantial amounts of organic carbon (OC) and nutrients which have accumulated in late Pleistocene and Holocene unconsolidated deposits. Permafrost vulnerability to thaw subsidence, collapsing coastlines and irreversible landscape change are largely due to the presence of large amounts of massive ground ice such as ice wedges. However, ground ice has not, until now, been considered to be a source of dissolved organic carbon (DOC), dissolved inorganic carbon (DIC) and other elements which are important for ecosystems and carbon cycling. Here we show, using biogeochemical data from a large number of different ice bodies throughout the Arctic, that ice wedges have the greatest potential for DOC storage, with a maximum of 28.6 mg L-1 (mean: 9.6 mg L-1). Variation in DOC concentration is positively correlated with and explained by the concentrations and relative amounts of typically terrestrial cations such as Mg2+ and K+. DOC sequestration into ground ice was more effective during the late Pleistocene than during the Holocene, which can be explained by rapid sediment and OC accumulation, the prevalence of more easily degradable vegetation and immediate incorporation into permafrost. We assume that pristine snowmelt is able to leach considerable amounts of well-preserved and highly bioavailable DOC as well as other elements from surface sediments, which are rapidly frozen and stored in ground ice, especially in ice wedges, even before further degradation. We found that ice wedges in the Yedoma region represent a significant DOC (45.2 Tg) and DIC (33.6 Tg) pool in permafrost areas and a freshwater reservoir of 4200 km(2). This study underlines the need to discriminate between particulate OC and DOC to assess the availability and vulnerability of the permafrost car-bon pool for ecosystems and climate feedback upon mobilization.