Einar Heegaard

Comparing palaeolimnological and instrumental evidence of climate change for remote mountain lakes over the last 200 years

This paper compares the palaeolimnological evidence for climate change over the last 200 years with instrumental climate data for the same period at seven European remote mountain lakes. The sites are Øvre Neådalsvatn (Norway), Saanajärvi (Finland), Gossenköllesee (Austria), Hagelseewli (Switzerland), Jezero v Ledvici (Slovenia), Estany Redó (Spain, Pyrenees), and Nižné Terianske Pleso (Slovakia). We used multiple regression analysis to transfer homogenised lowland air temperature records to each of the sites, and these reconstructions were validated using data from on-site automatic weather stations. These data showed that mean annual temperature has varied over the last 200 years at each site by between 1 and 2 °C, typical of the high frequency variability found throughout the Holocene, and appropriate, therefore, to test the sensitivity of the various proxy methods used. Sediment cores from each site were radiometrically dated using 210Pb, 137Cs and 241Am and analysed for loss-on-ignition, C, N, S, pigments, diatoms, chrysophytes, Cladocera and chironomids. Comparisons between the proxy data and the instrumental data were based on linear regression analysis with the proxy data treated as response variables and the instrumental data (after smoothing using LOESS regressions) as predictor variables. The results showed few clear or consistent patterns with generally low or very low r2 values. Highest values were found when the data were compared after smoothing using a broad span, indicating that some of the proxy data were capturing climate variability but only at a relatively coarse time resolution. Probable reasons for the weak performance of the methods used include inaccurate dating, especially for earlier time periods, the influence of confounding forcing factors at some sites e.g., air pollution, earthquakes, and the insensitivity of some methods to low amplitude climate forcing. Nevertheless, there were trends in some proxy records at a number of sites that had a relatively unambiguous correspondence with the instrumental climate records. These included organic matter and associated variables (C and N) and planktonic diatom assemblages at the majority of sites and chrysophytes and chironomids at a few sites. Overall for longer term studies of the Holocene, these results indicate the need to be cautious in the interpretation of proxy records, the importance of proxy method validation, the continuing need to use reinforcing multi-proxy approaches, and the need for careful site and method selection.

Species-environmental relationships of aquatic macrophytes in Northern Ireland

Abstract A data-set from 574 lakes in Northern Ireland was used to analyse the individual relationships of the occurrence of aquatic macrophytes to the lake environment. The 32 most frequent species were used in the analyses. The 21 environmental parameters recorded for each lake were divided into five groups; (1) geographical features; (2) lake physical variables; (3) lake-water chemistry summary variables; (4) lake-water major ions; and (5) lake-water nutrients. The species–environmental relationships were analysed using generalised additive models (GAM) and canonical correspondence analysis (CCA). The results were interpreted in terms of expected range, tolerance, and optimum of each species. Predictive models were made for each of the most frequent aquatic macrophytes along individual environmental gradients. The most influential variables are related to local-scale chemical and nutrient composition, both of which are highly correlated with altitude because hardwater, nutrient-rich lakes are restricted to the lowlands. Major ions and nutrients constrain the species into two groups associated with either low or high ionic concentration. The high ionic concentration group of species may be further divided into those associated with calcareous lakes or lakes with high sodium and chloride. Local-scale variation in lake chemistry and the strong environmental responses of the aquatic macrophytes suggest a strong local influence on species composition of different lakes, leading to the interpretation that the occurrence of a species in a lake is predominantly controlled by the catchment use, particularly farming and fertiliser use.

Climate change and spring-fruiting fungi

Most macrofungi produce ephemeral fruit bodies during autumn but some have adapted to spring fruiting. In this study, temporal changes in the time of spring fruiting in Norway and the UK during 1960–2007 have been investigated by statistical analyses of about 6000 herbarium and field records, covering 34 species. Nearly 30 per cent of the temporal variation in fruiting could be ascribed to spatial and species-specific effects. Correcting for these effects, linear trends towards progressively earlier fruiting were detected during the entire period in both Norway and the UK, with a change in average fruiting day of 18 days over the study period. Early fruiting was correlated with high winter temperatures in both countries, indicating that the observed phenological changes are likely due to earlier onset of spring. There were also significant correlations between climatic conditions in one year and timing of fruiting the following year, indicating that below-ground mycelia are influenced by climatic conditions over a longer time period before fruiting. Fruiting dates were, however, not strictly related to changes in vernal accumulated thermal time. Our results indicate that global warming has lead to progressively earlier fruiting of spring fungi in northwest Europe during the last half century.

Warming-induced shift in European mushroom fruiting phenology

In terrestrial ecosystems, fungi are the major agents of decomposition processes and nutrient cycling and of plant nutrient uptake. Hence, they have a vital impact on ecosystem processes and the terrestrial carbon cycle. Changes in productivity and phenology of fungal fruit bodies can give clues to changes in fungal activity, but understanding these changes in relation to a changing climate is a pending challenge among ecologists. Here we report on phenological changes in fungal fruiting in Europe over the past four decades. Analyses of 746,297 dated and geo-referenced mushroom records of 486 autumnal fruiting species from Austria, Norway, Switzerland, and the United Kingdom revealed a widening of the annual fruiting season in all countries during the period 1970–2007. The mean annual day of fruiting has become later in all countries. However, the interspecific variation in phenological responses was high. Most species moved toward a later ending of their annual fruiting period, a trend that was particularly strong in the United Kingdom, which may reflect regional variation in climate change and its effects. Fruiting of both saprotrophic and mycorrhizal fungi now continues later in the year, but mycorrhizal fungi generally have a more compressed season than saprotrophs. This difference is probably due to the fruiting of mycorrhizal fungi partly depending on cues from the host plant. Extension of the European fungal fruiting season parallels an extended vegetation season in Europe. Changes in fruiting phenology imply changes in mycelia activity, with implications for ecosystem function.

Seasonal trends in the biomass and structure of bryophyte-associated fungal communities explored by 454 pyrosequencing

Bryophytes are a dominant vegetation component of the boreal forest, but little is known about their associated fungal communities, including seasonal variation within them. Seasonal variation in the fungal biomass and composition of fungal communities associated with three widespread boreal bryophytes was investigated using HPLC assays of ergosterol and amplicon pyrosequencing of the internal transcribed spacer 2 (ITS2) region of rDNA. The bryophyte phyllosphere community was dominated by Ascomycota. Fungal biomass did not decline appreciably in winter (P=0.272). Significant host-specific patterns in seasonal variation of biomass were detected (P=0.003). Although seasonal effects were not the primary factors structuring community composition, collection date significantly explained (P=0.001) variation not attributed to locality, host, and tissue. Community homogenization and a reduction in turnover occurred with the onset of frost events and subzero air and soil temperatures. Fluctuations in the relative abundance of particular fungal groups seem to reflect the nature of their association with mosses, although conclusions are drawn with caution because of potential methodological bias. The moss-associated fungal community is dynamic, exhibiting seasonal turnover in composition and relative abundance of different fungal groups, and significant fungal biomass is present year-round, suggesting a winter-active fungal community.

The outer border and central border for species–environmental relationships estimated by non-parametric generalised additive models

Abstract In modelling species–environmental relationships, non-parametric generalised additive modelling (GAM) regression has become a popular approach, as it has been shown that the generalised linear model (GLM) approach includes a bias that may be biologically unrealistic. However, the more versatile GAM approach has the disadvantage that a simple numerical description of the estimated response and estimation of the range (2×tolerance) and tolerance ( t = σ , sigma) are not possible. To try to overcome this, I introduce a procedure that estimates the outer border and central border numerically for non-parametric functions. The outer and central borders are given by the distance from the optimum that is needed for the response curve to drop a certain amount, i.e. these parameters represent the rate of decrease in the response in both directions from the optimum independent of each other. Thus, the species with a narrow niche will show a faster drop (shorter distance from the optimum) than the species with a broader niche. The outer and the central borders are defined as the predictor values which take the response value equal to a specified fraction of the maximum response: max( E ( y ∣ x ))×exp(−2) for the outer border and max( E ( y ∣ x ))×exp(−1/2) for the expected central border. These parameters are directly related to the estimated response values rather than, as both range and tolerance, related to a polynomial function defining the response curve. For logistic regressions there are increasing number of positive observations outside the classic range as the maximum response increases. In this, the expected borders differ from the range, as the interpretation of the former is independent of the maximum response. By reporting the lower and upper outer border and the central border combined with the optimum for the species, a simple unbiased numerical description of the species–environment response is obtained, which can be used in comparative studies.

Estimation of Age-Depth Relationships

An accurate and precise chronology is an essential pre-requisite for any palaeolimnological study. Chronologies give time-scales for events, and hence for rates for patterns and processes, and make it possible to compare and correlate events in different stratigraphical sequences. Palaeolimnology without chronology is history without dates.

Scale-dependent importance of environment, land use and landscape structure for species richness and composition of SE Norwegian modern agricultural landscapes

Knowledge of variation in vascular plant species richness and species composition in modern agricultural landscapes is important for appropriate biodiversity management. From species lists for 2201 land-type patches in 16 1-km2 plots five data sets differing in sampling-unit size from patch to plot were prepared. Variation in each data set was partitioned into seven sources: patch geometry, patch type, geographic location, plot affiliation, habitat diversity, ecological factors, and land-use intensity. Patch species richness was highly predictable (75% of variance explained) by patch area, within-patch heterogeneity and patch type. Plot species richness was, however, not predictable by any explanatory variable, most likely because all studied landscapes contained all main patch types – ploughed land, woodland, grassland and other open land – and hence had a large core of common species. Patch species composition was explained by variation along major environmental complex gradients but appeared nested to lower degrees in modern than in traditional agricultural landscapes because species-poor parts of the landscape do not contain well-defined subsets of the species pool of species-rich parts. Variation in species composition was scale dependent because the relative importance of specific complex gradients changed with increasing sampling-unit size, and because the amount of randomness in data sets decreased with increasing sampling-unit size. Our results indicate that broad landscape structural changes will have consequences for landscape-scale species richness that are hard or impossible to predict by simple surrogate variables.

Mushroom's spore size and time of fruiting are strongly related: is moisture important?

Most basidiomycete fungi produce annual short-lived sexual fruit bodies from which billions of microscopic spores are spread into the air during a short time period. However, little is known about the selective forces that have resulted in some species fruiting early and others later in the fruiting season. This study of relationships between morphological and ecological characteristics, climate factors and time of fruiting are based upon thorough statistical analyses of 66 520 mapped records from Norway, representing 271 species of autumnal fruiting mushroom species. We found a strong relationship between spore size and time of fruiting; on average, a doubling of spore size (volume) corresponded to 3 days earlier fruiting. Small-spored species dominate in the oceanic parts of Norway, whereas large-spored species are typical of more continental parts. In separate analyses, significant relationships were observed between spore size and climate factors. We hypothesize that these relationships are owing to water balance optimization, driven by water storage in spores as a critical factor for successful germination of primary mycelia in the drier micro-environments found earlier in the fruiting season and/or in continental climates.

Amplicon-pyrosequencing-based detection of compositional shifts in bryophyte-associated fungal communities along an elevation gradient

Although bryophytes are a dominant vegetation component of boreal and alpine ecosystems, little is known about their associated fungal communities. HPLC assays of ergosterol (fungal biomass) and amplicon pyrosequencing of the ITS2 region of rDNA were used to investigate how the fungal communities associated with four bryophyte species changed across an elevational gradient transitioning from conifer forest to the low‐alpine. Fungal biomass and OTU richness associated with the four moss hosts did not vary significantly across the gradient (P > 0.05), and both were more strongly affected by host and tissue type. Despite largely constant levels of fungal biomass, distinct shifts in community composition of fungi associated with Hylocomium, Pleurozium and Polytrichum occurred between the elevation zones of the gradient. This likely is a result of influence on fungal communities by major environmental factors such as temperature, directly or indirectly mediated by, or interacting with, the response of other components of the vegetation (i.e. the dominant trees). Fungal communities associated with Dicranum were an exception, exhibiting spatial autocorrelation between plots, and no significant structuring by elevation. Nevertheless, the detection of distinct fungal assemblages associated with a single host growing in different elevation zones along an elevational gradient is of particular relevance in the light of the ongoing changes in vegetation patterns in boreal and alpine systems due to global climate warming.