H. J. B. Birks

Recent warming reverses long-term arctic cooling.

Climate Reversal The climate and environment of the Arctic have changed drastically over the short course of modern observation. Kaufman et al. (p. 1236) synthesized 2000 years of proxy data from lakes above 60° N latitude with complementary ice core and tree ring records, to create a paleoclimate reconstruction for the Arctic with a 10-year resolution. A gradual cooling trend at the start of the record had reversed by the beginning of the 20th century, when temperatures began to increase rapidly. The long-term cooling of the Arctic is consistent with a reduction in summer solar insolation caused by changes in Earths orbit, while the rapid and large warming of the past century is consistent with the human-caused warming. A 2000-year-long Arctic cooling trend seen in a surface air temperature reconstruction was reversed during the last century. The temperature history of the first millennium C.E. is sparsely documented, especially in the Arctic. We present a synthesis of decadally resolved proxy temperature records from poleward of 60°N covering the past 2000 years, which indicates that a pervasive cooling in progress 2000 years ago continued through the Middle Ages and into the Little Ice Age. A 2000-year transient climate simulation with the Community Climate System Model shows the same temperature sensitivity to changes in insolation as does our proxy reconstruction, supporting the inference that this long-term trend was caused by the steady orbitally driven reduction in summer insolation. The cooling trend was reversed during the 20th century, with four of the five warmest decades of our 2000-year-long reconstruction occurring between 1950 and 2000.

D.G. Frey and E.S. Deevey Review 1: Numerical tools in palaeolimnology – Progress, potentialities, and problems

In the last decade, palaeolimnology has shifted emphasis from being a predominantly qualitative, descriptive subject to being a quantitative, analytical science with the potential to address critical hypotheses concerning the impacts of environmental changes on limnic systems. This change has occurred because of (1) major developments in applied statistics, some of which have only become possible because of the extraordinary developments in computer technology, (2) increased concern about problem definition, research hypotheses, and project design, (3) the building up of high quality modern calibration data-sets, and (4) the narrowing of temporal resolution in palaeolimnological studies from centuries to decades or even single years or individual seasons.The most significant development in quantitative palaeolimnology has been the creation of many modern calibration data-sets of biotic assemblages and associated environmental data. Such calibration sets, when analysed by appropriate numerical procedures, have the potential to transform fossil biostratigraphical data into quantitative estimates of the past environment. The relevant numerical techniques are now well developed, widely tested, and perform remarkably well. The properties of these techniques are becoming better known as a result of simulation studies. The advantages and disadvantages of the preferred technique (weighted averaging partial least squares) are reviewed and the problems in model selection are discussed. The need for evaluation and validation of reconstructions is emphasised. Several statistical surprises have emerged from calibration studies. Outstanding problems remain the need for a detailed and consistent taxonomy in the calibration sets, the quality, representativeness, and inherent variability of the environmental variables of interest, and the inherent bias in the calibration models. Besides biological- environmental calibration sets, there is the potential to develop modern sediment-environment calibration sets to link sedimentary properties to catchment parameters. The adoption of a ‘dynamic calibration set’ approach may help to minimise the inherent bias in current calibration models. Modern regression techniques are available to explore the vast amount of unique ecological information about taxon-environment relationships in calibration data-sets.Hypothesis testing in palaeolimnology can be attempted directly by careful project design to take account of ‘natural experiments’ or indirectly by means of statistical testing, often involving computer- intensive permutation tests to evaluate specific null hypotheses. The validity of such tests depends on the type of permutation used in relation to the particular data-set being analysed, the sampling design, and the research questions being asked. Stratigraphical data require specific permutation tests. Several problems remain unsolved in devising permutation designs for fine-resolution stratigraphical data and for combined spatial and temporal data. Constrained linear or non-linear reduced rank regression techniques (e.g. redundancy analysis, canonical correspondence analysis and their partial counterparts) provide powerful tools for testing hypotheses in palaeolimnology. Work is needed, however, to extend their use to investigate and test for lag responses between biological assemblages and their environment.Having developed modern calibration data-sets, many palaeolimnologists are returning to the sedimentary record and are studying stratigraphical changes. In contrast to much palynological data, palaeolimnological data are often fine-resolution and as a result are often noisy, large, and diverse. Recent developments for detecting and summarising patterns in such data are reviewed, including statistical evaluation of zones, summarisation by detrended correspondence analysis, and non-parametric regression (e.g. LOESS). Techniques of time-series analysis that are free of many of the assumptions of conventional time-series analysis due to the development of permutation tests to assess statistical significance are of considerable potential in analysing fine-resolution palaeolimnological data. Such data also contain a wealth of palaeopopulation information. Robust statistical techniques are needed to help identify non-linear deterministic dynamics (chaos) from noise or random effects in fine-resolution palaeolimnological data.

What is natural? The need for a long-term perspective in biodiversity conservation.

Ecosystems change in response to factors such as climate variability, invasions, and wildfires. Most records used to assess such change are based on short-term ecological data or satellite imagery spanning only a few decades. In many instances it is impossible to disentangle natural variability from other, potentially significant trends in these records, partly because of their short time scale. We summarize recent studies that show how paleoecological records can be used to provide a longer temporal perspective to address specific conservation issues relating to biological invasions, wildfires, climate change, and determination of natural variability. The use of such records can reduce much of the uncertainty surrounding the question of what is “natural” and thereby start to provide important guidance for long-term management and conservation.

The use of Rarefaction Analysis for Estimating Palynological Richness from Quaternary Pollen-Analytical Data

Reconstructing temporal changes in diversity from pollen assemblages is potentially important both palaeoecologically and ecologically because community diversity may, in part, result from historical processes. The use of diversity indices such as Shannons information index or Simpsons index is not appropriate with pollen percentage data because such indices consider both the numbers of different taxa and their relative frequencies or representation. The latter aspect in pollen data is inevitably influenced by inherent differences in pollen production and dispersal. The total number of taxa present in a sample is a robust and useful measure of palynological richness if, and only if, all the pollen counts are standardized to a fixed number of grains. Rarefaction analysis implements such a standardization and provides minimum variance unbiased estimates of the expected number of taxa (t) in a random sample of n individuals taken from a larger collection of N individuals containing T taxa. The underlying mathematical theory of rarefaction analysis and its important biological and palaeoecological assumptions are discussed. The use of rarefaction analysis is illustrated with three data-sets: Crose Mere, central England (0-c. 12 500 BP); Abernethy Forest, eastern Scotland (5500-12 100 BP); and three sites (Lochs Ashik, Cleat, Meodal) on the Isle of Skye, western Scotland, all covering the last 10 500 years. Palynological richness, as estimated by rarefaction analysis, is high in the protocratic phase (c. 9500-12 500 BP), low in the mesocratic phase (c. 5500-9500 BP), low in the oligocratic phase (0-c. 5500 BP), and high in the Homo sapiens phase (0-c. 5000 BP) of the Holocene. Although factors such as local site characteristics and pollen production, dispersal, and input may influence temporal changes in richness, changes in palynological richness are interpreted as reflecting predominantly the changing floristic richness of the vegetation types in the pollen-source area of a lake and the changing mosaic structure of the landscape through time. Intermediate levels of disturbance, either natural in the protocratic phase or anthropogenic in the Homo sapiens phase appear to be important in maximizing richness at the landscape scale by preventing (he dominance of any single component but insufficient to cause extinction of components at the landscape scale.

July mean temperature and annual precipitation trends during the Holocene in the Fennoscandian tree-line area: pollen-based climate reconstructions

July mean temperature and annual precipipation during the last 9900 cal. yr BP were recon structed from pollen assemblages preserved in a sediment core from northern Finland. Quantitative recon structions were performed using a modern pollen-climate calibration model based on weighted-averaging partial least squares regression. The predictive ability of the model was evaluated against modern meteoro logical data using leave-one-out cross-validation. The prediction error for July mean temperature is c.1.0°C and for annual precipitation 340 mm. The July mean temperatures during the earliest Holocene were low, c.11.0°C, and annual precipitation was high, c. 600–800 mm. Between 8200 and 6700 cal. yr BP July mean temperatures reached their maxima, 12.5–13.0°C, which are c. 1.4–1.7°C higher than at present. At the same time precipitation decreased. During the late Holocene, July mean temperatures declined and the last 2000 years have been the coolest since the early Holocene. Precipitation has slightly increased. The spatial coherence between our results and of several other climate reconstructions from northern Europe indicates that the Holocene climate was strongly influenced by North Atlantic oceanic and atmospheric circulation patterns. We propose that the distinctly oceanic climate of the early Holocene was due to enhanced westerly (latitudinal) airflow which was replaced at c. 8200 cal. yr BP by a more meridional flow pattern and by the development of predominantly anticyclonic summer conditions.

Recent increases in species richness and shifts in altitudinal distributions of Norwegian mountain plants

Opportunities for observing long-term changes in natural biota are rare. Observations on the distribution and frequency of vascular plants were performed on 23 mountains situated along a west–east gradient in Jotunhei men, central Norway, where detailed site descriptions and species lists exist from ad 1930–31. The sites were resurveyed during the summer of 1998, to examine possible changes in species richness and species distributions along the altitudinal gradient during a 68-year period. Increased species richness was found on 19 of the mountains and was most pronounced at lower altitudes and in the eastern areas. Lowland species, dwarf shrubs and species with wide altitudinal and ecological ranges showed the greatest increases in abundance and altitudinal advances since the 1930–31 study. Species with more restricted habitat demands, such as some hygrophilous snow-bed species, have declined. High-altitude species have disappeared from their lower-elevation sites and increased their abundance at the highest altitudes. Climatic warming occurring in the last 100 years might have allowed the invasion of lowland and lee-slope species. Increased competition at sites where such species have invaded may have led to a decreased abundance of the less competitive species and a concentration of high-altitude species on the highest ridges. Natural succession since the ‘Little Ice Age’, increased deposition of nitrogen during recent years and changes in grazing and tourism might have in‘ uenced some of the species turnovers, but recent climatic changes are considered to be the most likely major driving factor for the changes observed.

The development of the aquatic ecosystem at Kråkenes Lake, western Norway, during the late glacial and early Holocene - a synthesis

This paper synthesises the palaeoecological reconstructions, including palaeoclimatic inferences, based on the available fossil record of plants (pollen, macrofossils, mosses, diatoms) and animals (chironomids, Cladocera, Coleoptera, Trichoptera, oribatid mites) in the late-glacial and early-Holocene sediments of Kråkenes Lake, western Norway, with special emphasis on changes in the aquatic ecosystem. New percentage and influx pollen diagrams for selected taxa provide insights into the terrestrial setting. The information from all the proxies is collated in a stratigraphical chart, and the inferred changes in the lake and its catchment are discussed. The individual fossil sequences are summarised by detrended correspondence analysis (DCA), and sample scores on the first DCA axes are plotted against an estimated calendar-year timescale for comparison of the timing and magnitude of changes in assemblage composition. The DCA plots show that the large late-glacial biotic changes were synchronous, and were driven by the overriding forcing factor of temperature. During the early Holocene, however, the changes in different groups were more gradual and were independent of each other, showing that other factors were important and interactive, such as the inwash of dissolved and particulate material from the catchment, the base and nutrient status of the lake-water, and the internal processes of ecosystem succession and sediment accumulation. This multi-disciplinary study, with proxies for changes in the lake and in the catchment, highlights the dependence of lake biota and processes not only on regional climatic changes but also on changes in the lake catchment and on internal processes within the lake. Rates of change for each group are also estimated and compared. The reaction times to the sharp temperature changes at the start and end of the Younger Dryas were very rapid and occurred within a decade of the temperature change. Aquatic organisms tracked the temperature and environmental changes very closely, and are probably the best recorders of late-glacial climatic change in the fossil record.

WACALIB version 3.3 — a computer program to reconstruct environmental variables from fossil assemblages by weighted averaging and to derive sample-specific errors of prediction

A computer program for reconstructing environmental variables (e.g. lake-water pH) from fossil assemblages (e.g. diatoms) by weighted averaging regression and calibration is described. The estimation of sample-specific errors of prediction by bootstrapping is outlined. The program runs on IBM-compatible personal computers.

Chironomid-inferred late-glacial and early-Holocene mean July air temperatures for Kråkenes Lake, western Norway

A chironomid data-set calibrated to July air temperatures, based on 44 lakes in western Norway, is used to reconstruct mean July air temperatures from late-glacial and early-Holocene fossil chironomid assemblages at Kråkenes Lake. The calibration function is based on Weighted Averaging Partial Least Squares regression and has a root mean square error of prediction (RMSEP) of 1.13 °C, a r2 of 0.69, and a maximum bias of 2.66 °C. All these statistics are based on leave-one-out cross-validation. A calibration function based on summer surface-water temperatures has a poorer performance (RMSEP = 2.22 °C, r2 = 0.30, maximum bias = 5.29 °C). The reconstructed July air temperatures at Kråkenes rise to 10.5 °C soon after deglaciation, are about 11.5 °C in the Allerød, decrease to 9.5-10 °C in the Younger Dryas, and rise rapidly within 15 yrs to 11.5 °C at the onset of the Holocene. There is a two-step rise to 13 °C or more in the early-Holocene. The likely over-estimation of Younger Dryas temperatures and under-estimation of early-Holocene temperatures probably result from the limited temperature range represented by the existing calibration set. The data set is currently being expanded to include lakes with warmer air temperatures (> 14 °C) and with colder air temperatures (< 8 °C).

Quantification of biotic responses to rapid climatic changes around the Younger Dryas — a synthesis

To assess the presence or absence of lags in biotic responses to rapid climatic changes, we: (1) assume that the δ18O in biogenically precipitated carbonates record global or hemispheric climatic change at the beginning and at the end of the Younger Dryas without any lag at our two study sites of Gerzensee and Leysin, Switzerland; (2) derive a time scale by correlating the δ18O record from these two sites with the δ18O record of the GRIP ice core; (3) measure δ18O records in ostracods and molluscs to check the record in the bulk samples and to detect possible hydrological changes; (4) analyse at Gerzensee and Leysin as well as at two additional sites (that lack carbonates and hence a δ18O record) pollen, plant macrofossils, chironomids, beetles and other insects, and Cladocera; (5) estimate our sampling resolution using the GRIP time scale for the isotope stratigraphies and the biostratigraphies; and (6) summarise the major patterns of compositional change in the biostratigraphies by principal component analysis or correspondence analysis. We conclude that, at the major climatic shifts at the beginning and end of the Younger Dryas, hardly any biotic lags occur (within the sampling resolution of 8–30 years) and that upland vegetation responded as fast as aquatic invertebrates. We suggest that the minor climatic changes associated with the Gerzensee and Preboreal oscillations were weakly recorded in the biostratigraphies at the lowland site, but were more distinct at higher altitudes. Individualistic responses of plant and animal species to climatic change may reflect processes in individuals (e.g. productivity and phenology), in populations (e.g. population dynamics), in spatial distributions (e.g. migrations), and in ecosystems (e.g. trophic state). We suggest that biotic responses may be telescoped together into relatively short periods (50 to 150 years), perhaps disrupting functional interactions among species and thus destabilising ecosystems.