Results of n-alkane biomarker analyses and paleoclimate reconstruction

The current study presents quantitative reconstructions of tree cover, annual precipitation and mean July temperature derived from the pollen record from Lake Billyakh (65°17'N, 126°47'E, 340 m above sea level) spanning the last ca. 50 kyr. The reconstruction of tree cover suggests presence of woody plants through the entire analyzed time interval, although trees played only a minor role in the vegetation around Lake Billyakh prior to 14 kyr BP (<5%). This result corroborates low percentages of tree pollen and low scores of the cold deciduous forest biome in the PG1755 record from Lake Billyakh. The reconstructed values of the mean temperature of the warmest month ~8-10 °C do not support larch forest or woodland around Lake Billyakh during the coldest phase of the last glacial between ~32 and ~15 kyr BP. However, modern cases from northern Siberia, ca. 750 km north of Lake Billyakh, demonstrate that individual larch plants can grow within shrub and grass tundra landscape in very low mean July temperatures of about 8 °C. This makes plausible our hypothesis that the western and southern foreland of the Verkhoyansk Mountains could provide enough moist and warm microhabitats and allow individual larch specimens to survive climatic extremes of the last glacial. Reconstructed mean values of precipitation are about 270 mm/yr during the last glacial interval. This value is almost 100 mm higher than modern averages reported for the extreme-continental north-eastern Siberia east of Lake Billyakh, where larch-dominated cold deciduous forest grows at present. This suggests that last glacial environments around Lake Billyakh were never too dry for larch to grow and that the summer warmth was the main factor, which limited tree growth during the last glacial interval. The n-alkane analysis of the Siberian plants presented in this study demonstrates rather complex alkane distribution patterns, which challenge the interpretation of the fossil records. In particular, extremely low n-alkane concentrations in the leaves of local coniferous trees and shrubs suggest that their contribution to the litter and therefore to the fossil lake sediments might be not high enough for tracing the Quaternary history of the needleleaved taxa using the n-alkane biomarker method.

Plant species, biomass and environmental characteristics of relevés along the North America Arctic bioclimate gradient

Question: How do interactions between the physical environment and biotic properties of vegetation influence the formation of small patterned-ground features along the Arctic bioclimate gradient? Location: At 68° to 78°N: six locations along the Dalton Highway in arctic Alaska and three in Canada (Banks Island, Prince Patrick Island and Ellef Ringnes Island). Methods: We analysed floristic and structural vegetation, biomass and abiotic data (soil chemical and physical parameters, the n-factor [a soil thermal index] and spectral information [NDVI, LAI]) on 147 microhabitat releves of zonalpatterned-ground features. Using mapping, table analysis (JUICE) and ordination techniques (NMDS). Results: Table analysis using JUICE and the phi-coefficient to identify diagnostic species revealed clear groups of diagnostic plant taxa in four of the five zonal vegetation complexes. Plant communities and zonal complexes were generally well separated in the NMDS ordination. The Alaska and Canada communities were spatially separated in the ordination because of different glacial histories and location in separate floristic provinces, but there was no single controlling environmental gradient. Vegetation structure, particularly that of bryophytes and total biomass, strongly affected thermal properties of the soils. Patterned-ground complexes with the largest thermal differential between the patterned-ground features and the surrounding vegetation exhibited the clearest patterned-ground morphologies.

(Table A1) Annual biomass of plants and animals recorded on Bylot Island between 1993-2009

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator--rey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.

Stable hydrogen and carbon isotope records and concentration data for four long chain n-alkanes from core GIK16160-3 derived from the Zambezi delta in the Mozambique Channel during the last 37,000 years

Hydrogen isotope values (dD) of sedimentary terrestrial leaf wax such as n-alkanes or n-acids have been used to map and understand past changes in rainfall amount in the tropics because dD of precipitation is commonly assumed as the first order controlling factor of leaf wax dD. Plant functional types and their photosynthetic pathways can also affect leaf wax dD but these biological effects are rarely taken into account in paleo studies relying on this rainfall proxy. To investigate how biological effects may influence dD values we here present a 37,000-year old record of dD and stable carbon isotopes (d13C) measured on four n-alkanes (n-C27, n-C29, n-C31, n-C33) from a marine sediment core collected off the Zambezi River mouth. Our paleo d13C records suggest that each individual n-alkanes had different C3/C4 proportional contributions. n-C29 was mostly derived from a C3 dicots (trees, shrubs and forbs) dominant vegetation throughout the entire record. In contrast, the longer chain n-C33 and n-C31 were mostly contributed by C4 grasses during the Glacial period but shifted to a mixture of C4 grasses and C3 dicots during the Holocene. Strong correlations between dD and d13C values of n-C33 (correlation coefficient R2 = 0.75, n = 58) and n-C31 (R2 = 0.48, n = 58) suggest that their dD values were strongly influenced by changes in the relative contributions of C3/C4 plant types in contrast to n-C29 (R2 = 0.07, n = 58). Within regions with variable C3/C4 input, we conclude that dD values of n-C29 are the most reliable and unbiased indicator for past changes in rainfall, and that dD and d13C values of n-C31 and n-C33 are sensitive to C3/C4 vegetation changes. Our results demonstrate that a robust interpretation of palaeohydrological data using n-alkane dD requires additional knowledge of regional vegetation changes from which nalkanes are synthesized, and that the combination of dD and d13C values of multiple n-alkanes can help to differentiate biological effects from those related to the hydrological cycle.

(Table 2) Radiocarbon ages of lake sediments from Boresø, East Greenland

This chapter provides a review of proxy data from a variety of natural archives sampled in the Wollaston Forland region, central Northeast Greenland. The data are used to describe long-term environmental and climatic changes. The focus is on reconstructing the Holocene conditions particularly in the Zackenberg area. In addition, this chapter provides an overview of the archaeological evidence for prehistoric occupation of the region. The Zackenberg area has been covered by the Greenland Ice Sheet several times during the Quaternary. At the Last Glacial Maximum (LGM, about 22,000 years BP), temperatures were much lower than at present, and only very hardy organisms may have survived in the region, even if ice-free areas existed. Marked warming at around 11,700 years BP led to ice recession, and the Zackenberg area was deglaciated in the early Holocene, prior to 10,100 years BP. Rapid early Holocene land emergence was replaced by a slight transgression in the late Holocene. During the Holocene, summer solar insolation decreased in the north. Following deglaciation of the region, summer temperatures probably peaked in the early to mid-Holocene, as indicated by the occurrence of a southern beetle species. However, the timing for the onset of the Holocene thermal maximum is rather poorly constrained because of delayed immigration of key plant species. During the thermal maximum, the mean July temperature was at least 2-3°C higher than at present. Evidence for declining summer temperatures is seen at around 5500, 4500 and 3500 years BP. The cooling culminated during the Little Ice Age that peaked about 100-200 years ago. The first plants that immigrated to the region were herbs and mosses. The first dwarf shrubs arrived in Northeast Greenland prior to 10,400 years BP, and dwarf birch arrived around 8800 years BP. The first people arrived about 4500 years BP, but the region was depopulated several times before the last people disappeared some time after 1823 AD, perhaps as a consequence of poor hunting conditions during the peak of the Little Ice Age.

Paleoclimate reconstruction from Miocene macroflora in Kazakhstan compiled from various publications

25 datasets (13 fossil leaf and pollen assemblages, 12 quantitative palaeoclimatic datasets) are provided in order to analyse Early Miocene palaeoclimate in Kazakhstan. The rich fossil record in Kazakhstan documents that during the Oligocene and Early Miocene this area in Central Eurasia was densely forested with warm-temperate deciduous trees and shrubs of the so-called "Turgayan flora". 29 fossil floras from 13 localities have been selected for a quantitative analysis of the Aquitanian (early Early Miocene) climate situation in Kazakhstan. The assessed mean annual temperatures generally place around 15 °C, while values of mean annual precipitation are of about 1000 mm. In combination with several other climate parameters estimated (temperatures of warmest and coldest months, precipitation rates of wettest, driest and warmest months), these data reflect uniform climatic conditions over several thousands of square kilometres. Data of temperature parameters show slight spatial differentiations, with generally cooler mean annual temperatures and higher seasonality (i.e. warmer summers and colder winters) in the north-eastern part of the study area compared with the south-western area around Lake Aral. As compared with palaeoclimate estimates for the European and East Asian Aquitanian, the central part of the Eurasian continent reveals evident signals of higher seasonality and slightly increased continentality.

Multi-proxy records of Two-Yurts Lake

Within the scope of Russian-German palaeoenvironmental research, Two-Yurts Lake (TYL, Dvuh-Yurtochnoe in Russian) was chosen as the main scientific target area to decipher Holocene climate variability on Kamchatka. The 5x2 km large and 26 m deep lake is of proglacial origin and situated on the eastern flank of Sredinny Ridge at the northwestern end of the Central Kamchatka Valley, outside the direct influence of active volcanism. Here, we present results of a multi-proxy study on sediment cores, spanning about the last 7000 years. The general tenor of the TYL record is an increase in continentality and winter snow cover in conjunction with a decrease in temperature, humidity, and biological productivity after 5000-4500 cal yrs BP, inferred from pollen and diatom data and the isotopic composition of organic carbon. The TYL proxy data also show that the late Holocene was punctuated by two colder spells, roughly between 4500 and 3500 cal yrs BP and between 1000 and 200 cal yrs BP, as local expressions of the Neoglacial and Little Ice Age, respectively. These environmental changes can be regarded as direct and indirect responses to climate change, as also demonstrated by other records in the regional terrestrial and marine realm. Long-term climate deterioration was driven by decreasing insolation, while the short-term climate excursions are best explained by local climatic processes. The latter affect the configuration of atmospheric pressure systems that control the sources as well as the temperature and moisture of air masses reaching Kamchatka.

A 2.8 Ma sedimentological, geochemical and palynological record of sediment core ICDP5011-1 in Lake Elgygytgyn, NE Russia

The reliability of Arctic climate predictions is currently hampered by insufficient knowledge of natural climate variability in the past. A sediment core from Lake El'gygytgyn (NE Russia) provides a continuous high-resolution record from the Arctic spaning the past 2.8 Ma. The core reveals numerous "super interglacials" during the Quaternary, with maximum summer temperatures and annual precipitation during marine benthic isotope stages (MIS) 11c and 31 ~4-5 °C and ~300 mm higher than those of MIS 1 and 5e. Climate simulations show these extreme warm conditions are difficult to explain with greenhouse gas and astronomical forcing alone, implying the importance of amplifying feedbacks and far field influences. The timing of Arctic warming relative to West Antarctic Ice Sheet retreats implies strong interhemispheric climate connectivity.

A 3.6 Ma sedimentological, geochemical and palynological record of sediment core ICDP5011-1 in Lake Elgygytgyn, NE Russia

Understanding the evolution of Arctic polar climate from the protracted warmth of the middle Pliocene into the earliest glacial cycles in the Northern Hemisphere has been hindered by the lack of continuous, highly resolved Arctic time series. Evidence from Lake El'gygytgyn, NE Arctic Russia, shows that 3.6-3.4 million years ago, summer temperatures were ~8°C warmer than today when pCO2 was ~400 ppm. Multiproxy evidence suggests extreme warmth and polar amplification during the middle Pliocene, sudden stepped cooling events during the Pliocene-Pleistocene transition, and warmer than present Arctic summers until ~2.2 Ma, after the onset of Northern Hemispheric glaciation. Our data are consistent with sea-level records and other proxies indicating that Arctic cooling was insufficient to support large-scale ice sheets until the early Pleistocene.

Pollen data and pollen-based climate reconstruction for the KTK1 pollen record from Lake Kotokel

In this study a radiocarbon-dated pollen record from Lake Kotokel (52°47' N, 108°07' E, 458 m a.s.l.) located in southern Siberia east of Lake Baikal was used to derive quantitative characteristics of regional vegetation and climate from about 15 kyr BP (1 kyr = 1000 cal. yr) until today. Quantitative reconstruction of the late glacial vegetation and climate dynamics suggests that open steppe and tundra communities predominated in the study area prior to ca. 13.5 kyr BP and again during the Younger Dryas interval, between 12.8 and 11.6 kyr BP. The pollen-based climate reconstruction suggests lower-than-present mean January (~ -38 °C) and July (~ 12 °C) temperatures and annual precipitation (~ 270-300 mm) values during these time intervals. Boreal woodland replaced the primarily open landscape around Kotokel three times at about 14.8-14.7 kyr BP, during the Allerød Interstadial between 13.3-12.8 kyr BP and with the onset of the Holocene interglacial between 11.5 and 10.5 kyr BP, presumably in response to a noticeable increase in precipitation, and in July and January temperatures. The maximal spread of the boreal forest (taiga) communities in the region is associated with a warmer and wetter-than-present climate (Tw ~ 17-18 °C, Tc ~ -19 °C, Pann ~ 500-550 mm) that occurred ca. 10.8-7.3 kyr BP. During this time interval woody vegetation covered more than 50 % of the area within a 21x21 km window around the lake. The pollen-based best modern analogue reconstruction suggests a decrease in woody cover percentages and in all climatic variables about 7-6.5 kyr BP. Our results demonstrate a gradual decrease in precipitation and mean January temperature towards their present-day values in the region around Lake Kotokel since that time.

n-alkane carbon isotope ratios in plants and recent lake sediments from Cameroon

Trees and shrubs in tropical Africa use the C3 cycle as a carbon fixation pathway during photosynthesis, while grasses and sedges mostly use the C4 cycle. Leaf-wax lipids from sedimentary archives such as the long-chain n-alkanes (e.g., n-C27 to n-C33) inherit carbon isotope ratios that are representative of the carbon fixation pathway. Therefore, n-alkane d13C values are often used to reconstruct past C3/C4 composition of vegetation, assuming that the relative proportions of C3 and C4 leaf waxes reflect the relative proportions of C3 and C4 plants. We have compared the d13C values of n-alkanes from modern C3 and C4 plants with previously published values from recent lake sediments and provide a framework for estimating the fractional contribution (areal-based) of C3 vegetation cover (fC3) represented by these sedimentary archives. Samples were collected in Cameroon, across a latitudinal transect that accommodates a wide range of climate zones and vegetation types, as reflected in the progressive northward replacement of C3-dominated rain forest by C4-dominated savanna. The C3 plants analysed were characterised by substantially higher abundances of n-C29 alkanes and by substantially lower abundances of n-C33 alkanes than the C4 plants. Furthermore, the sedimentary d13C values of n-C29 and n-C31 alkanes from recent lake sediments in Cameroon (-37.4 per mil to -26.5 per mil) were generally within the range of d13C values for C3 plants, even when from sites where C4 plants dominated the catchment vegetation. In such cases simple linear mixing models fail to accurately reconstruct the relative proportions of C3 and C4 vegetation cover when using the d13C values of sedimentary n-alkanes, overestimating the proportion of C3 vegetation, likely as a consequence of the differences in plant wax production, preservation, transport, and/or deposition between C3 and C4 plants. We therefore tested a set of non-linear binary mixing models using d13C values from both C3 and C4 vegetation as end-members. The non-linear models included a sigmoid function (sine-squared) that describes small variations in the fC3 values as the minimum and maximum d13C values are approached, and a hyperbolic function that takes into account the differences between C3 and C4 plants discussed above. Model fitting and the estimation of uncertainties were completed using the Monte Carlo algorithm and can be improved by future data addition. Models that provided the best fit with the observed d13C values of sedimentary n-alkanes were either hyperbolic functions or a combination of hyperbolic and sine-squared functions. Such non-linear models may be used to convert d13C measurements on sedimentary n-alkanes directly into reconstructions of C3 vegetation cover.

Pollen records from Bol'shoy Lyakhovsky Island, Siberia

Cryolithological, ground ice and fossil bioindicator (pollen, diatoms, plant macrofossils, rhizopods, insects, mammal bones) records from Bol'shoy Lyakhovsky Island permafrost sequences (73°20'N, 141°30'E) document the environmental history in the region for the past c. 115 kyr. Vegetation similar to modern subarctic tundra communities prevailed during the Eemian/Early Weichselian transition with a climate warmer than the present. Sparse tundra-like vegetation and harsher climate conditions were predominant during the Early Weichselian. The Middle Weichselian deposits contain peat and peaty soil horizons with bioindicators documenting climate amelioration. Although dwarf willows grew in more protected places, tundra and steppe vegetation prevailed. Climate conditions became colder and drier c. 30 kyr BP. No sediments dated between c. 28.5 and 12.05 14C kyr BP were found, which may reflect active erosion during that time. Herb and shrubby vegetation were predominant 11.6-11.3 14C kyr BP. Summer temperatures were c. 4 °C higher than today. Typical arctic environments prevailed around 10.5 14C kyr BP. Shrub alder and dwarf birch tundra were predominant between c. 9 and 7.6 kyr BP. Reconstructed summer temperatures were at least 4 °C higher than present. However, insect remains reflect that steppe-like habitats existed until c. 8 kyr BP. After 7.6 kyr BP, shrubs gradually disappeared and the vegetation cover became similar to that of modern tundra. Pollen and beetles indicate a severe arctic environment c. 3.7 kyr BP. However, Betula nana, absent on the island today, was still present. Together with our previous study on Bol'shoy Lyakhovsky Island covering the period between about 200 and 115 kyr, a comprehensive terrestrial palaeoenvironmental data set from this area in western Beringia is now available for the past two glacial-interglacial cycles.