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The deciduous needle tree larch (Larix Mill.) covers more than 80% of the Asian boreal forests. Only a few Larix species constitute the vast forests and these species differ markedly in their ecological traits, most importantly in their ability to grow on and stabilize underlying permafrost. The pronounced dominance of the summergreen larches makes the Asian boreal forests unique, as the rest of the northern hemisphere boreal forests is almost exclusively dominated by evergreen needle-leaf forests. Global warming is impacting the whole world but is especially pronounced in the arctic and boreal regions. Although adapted to extreme climatic conditions, larch forests are sensitive to varying climatic conditions. By their sheer size, changes in Asian larch forests as range shifts or changes in species composition and the resulting vegetation-climate feedbacks are of global relevance. It is however still uncertain if larch forests will persist under the ongoing warming climate or if they will be replaced by evergreen forests. It is therefore of great importance to understand how these ecosystems will react to future climate warmings and if they will maintain their dominance. One step in the better understanding of larch dynamics is to study how the vast dominant forests developed and why they only established in northern Asia. A second step is to study how the species reacted to past changes in the climate.
The first objective of this thesis was to review and identify factors promoting Asian larch dominance. I achieved this by synthesizing and comparing reported larch occurrences and influencing components on the northern hemisphere continents in the present and in the past. The second objective was to find a possibility to directly study past Larix populations in Siberia and specifically their genetic variation, enabling the study of geographic movements. For this, I established chloroplast enrichment by hybridization capture from sedimentary ancient DNA (sedaDNA) isolated from lake sediment records. The third objective was to use the established method to track past larch populations, their glacial refugia during the Last Glacial Maximum (LGM) around 21,000 years before present (ka BP), and their post-glacial migration patterns.
To study larch promoting factors, I compared the present state of larch species ranges, areas of dominance, their bioclimatic niches, and the distribution on different extents and thaw depths of permafrost. The species comparison showed that the bioclimatic niches greatly overlap between the American and Asian species and that it is only in the extremely continental climates in which only the Asian larch species can persist. I revealed that the area of dominance is strongly connected to permafrost extent but less linked to permafrost seasonal thaw depths. Comparisons of the paleorecord of larch between the continents suggest differences in the recolonization history. Outside of northern Asia and Alaska, glacial refugial populations of larch were confined to the southern regions and thus recolonization could only occur as migration from south to north. Alaskan larch populations could not establish wide-range dominant forest which could be related to their own genetically depletion as separated refugial population. In Asia, it is still unclear whether or not the northern refugial populations contributed and enhanced the postglacial colonization or whether they were replaced by populations invading from the south in the course of climate warming. Asian larch dominance is thus promoted partly by adaptions to extremely continental climates and by adaptations to grow on continuous permafrost but could be also connected to differences in glacial survival and recolonization history of Larix species.
Except for extremely rare macrofossil findings of fossilized cones, traditional methods to study past vegetation are not able to distinguish between larch species or populations. Within the scope of this thesis, I therefore established a method to retrieve genetic information of past larch populations to distinguish between species. Using the Larix chloroplast genome as target, I successfully applied the method of DNA target enrichment by hybridization capture on sedaDNA samples from lake records and showed that it is able to distinguish between larch species. I then used the method on samples from lake records from across Siberia dating back up to 50 ka BP. The results allowed me to address the question of glacial survival and post-glacial recolonization mode in Siberian larch species. The analyzed pattern showed that LGM refugia were almost exclusively constituted by L. gmelinii, even in sites of current L. sibirica distribution. For included study sites, L. sibirica migrated into its extant northern distribution area only in the Holocene. Consequently, the post-glacial recolonization of L. sibirica was not enhanced by northern glacial refugia. In case of sites in extant distribution area of L. gmelinii, the absence of a genetic turn-over point to a continuous population rather than an invasion of southern refugia. The results suggest that climate has a strong influence on the distribution of Larix species and that species may also respond differently to future climate warming. Because species differ in their ecological characteristics, species distribution is also relevant with respect to further feedbacks between vegetation and climate.
With this thesis, I give an overview of present and past larch occurrences and evaluate which factors promote their dominance. Furthermore, I provide the tools to study past Larix species and give first important insights into the glacial history of Larix populations.
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.
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.
Climate change of anthropogenic origin is affecting Earth’s biodiversity and therefore ecosystems and their services. High latitude ecosystems are even more impacted than the rest of Northern Hemisphere because of the amplified polar warming. Still, it is challenging to predict the dynamics of high latitude ecosystems because of complex interaction between abiotic and biotic components. As the past is the key to the future, the interpretation of past ecological changes to better understand ongoing processes is possible. In the Quaternary, the Pleistocene experienced several glacial and interglacial stages that affected past ecosystems. During the last Glacial, the Pleistocene steppe-tundra was covering most of unglaciated northern hemisphere and disappeared in parallel to the megafauna’s extinction at the transition to the Holocene (~11,700 years ago). The origin of the steppe-tundra decline is not well understood and knowledge on the mechanisms, which caused shifts in past communities and ecosystems, is of high priority as they are likely comparable to those affecting modern ecosystems. Lake or permafrost core sediments can be retrieved to investigate past biodiversity at transitions between glacial and interglacial stages. Siberia and Beringia were the origin of dispersal of the steppe-tundra, which make investigation this area of high priority. Until recently, macrofossils and pollen were the most common approaches. They are designed to reconstruct past composition changes but have limit and biases. Since the end of the 20th century, sedimentary ancient DNA (sedaDNA) can also be investigated. My main objectives were, by using sedaDNA approaches to provide scientific evidence of compositional and diversity changes in the Northern Hemisphere ecosystems at the transition between Quaternary glacial and interglacial stages.
In this thesis, I provide snapshots of entire ancient ecosystems and describe compositional changes between Quaternary glacial and interglacial stages, and confirm the vegetation composition and the spatial and temporal boundaries of the Pleistocene steppe-tundra. I identify a general loss of plant diversity with extinction events happening in parallel of megafauna’ extinction. I demonstrate how loss of biotic resilience led to the collapse of a previously well-established system and discuss my results in regards to the ongoing climate change. With further work to constrain biases and limits, sedaDNA can be used in parallel or even replace the more established macrofossils and pollen approaches as my results support the robustness and potential of sedaDNA to answer new palaeoecological questions such as plant diversity changes, loss and provide snapshots of entire ancient biota.