Ericoid shrub encroachment shifts aboveground-belowground linkages in three peatlands across Europe and Western Siberia.

In northern peatlands, reduction of Sphagnum dominance in favour of vascular vegetation is likely to influence biogeochemical processes. Such vegetation changes occur as the water table lowers and temperatures rise. To test which of these factors has a significant influence on peatland vegetation, we conducted a 3-year manipulative field experiment in Linje mire (northern Poland). We manipulated the peatland water table level (wet, intermediate and dry; on average the depth of the water table was 17.4, 21.2 and 25.3 cm respectively), and we used open-top chambers (OTCs) to create warmer conditions (on average increase of 1.2°C in OTC plots compared to control plots). Peat drying through water table lowering at this local scale had a larger effect than OTC warming treatment per see on Sphagnum mosses and vascular plants. In particular, ericoid shrubs increased with a lower water table level, while Sphagnum decreased. Microclimatic measurements at the plot scale indicated that both water-level and temperature, represented by heating degree days (HDDs), can have significant effects on the vegetation. In a large-scale complementary vegetation gradient survey replicated in three peatlands positioned along a transitional oceanic-continental and temperate-boreal (subarctic) gradient (France-Poland-Western Siberia), an increase in ericoid shrubs was marked by an increase in phenols in peat pore water, resulting from higher phenol concentrations in vascular plant biomass. Our results suggest a shift in functioning from a mineral-N-driven to a fungi-mediated organic-N nutrient acquisition with shrub encroachment. Both ericoid shrub encroachment and higher mean annual temperature in the three sites triggered greater vascular plant biomass and consequently the dominance of decomposers (especially fungi), which led to a feeding community dominated by nematodes. This contributed to lower enzymatic multifunctionality. Our findings illustrate mechanisms by which plants influence ecosystem responses to climate change, through their effect on microbial trophic interactions.

Long-term microclimate study of a peatland in Central Europe to understand microrefugia.

Peatlands perform many important ecosystem functions at both the local and global scale, including hydrologic and climatic regulation. Although peatlands often act as climatic microrefugia, they have rarely been the subject of long-term microclimatic studies. In this study, we aimed to compare the local climatic conditions of a mid-forest mire to that of an open area and examine the differences in microclimates within the mire based on plant community diversity, shading, and water table depths. The peatland studied in this work was significantly cooler than the reference site, mainly due to a higher decline in nighttime air temperatures. However, the daily maximum air temperature near the ground was often higher. We also noticed that microclimates significantly differed within the studied peatland. Wet and shaded microsites were cooler than the sites having a lower water level and receiving higher amounts of solar radiation. The results of the study suggest that peatlands have locally cooler climates, and thus can serve as climate change refugia. These findings can help us interpret reconstructed data from the peat archive, and, when combined with experiments, identify tipping points for peatland ecosystems.

Unveiling tipping points in long-term ecological records from Sphagnum-dominated peatlands.

Unveiling past tipping points is a prerequisite for a better understanding of how individual species and entire ecosystems will respond to future climate change. Such knowledge is key for the implementation of biodiversity conservation. We identify the relationships between peatland vegetation and hydrological conditions over the past 2000 years using plant macrofossils, testate amoebae-based quantitative hydrological reconstructions and Sphagnum-moss functional traits from seven Polish peatland records. Using threshold indicator taxa analysis, we discovered that plant community composition strongly converged at a water level of ca 11.7 cm, indicating a community-level tipping point. We identified 45 plant taxa that showed either an increase or a decrease in their relative abundance between 8 and 17 cm of water-level depth. Our analysis of Sphagnum community traits further showed that Sphagnum functional diversity was remarkably stable over time despite Sphagnum species sensitivity to hydrological conditions. Our results suggest that past hydrological shifts did not influence major functions of the Sphagnum community, such as photosynthetic capacity, growth and productivity, owing to species replacement with a similar functional space. Although further studies including trait plasticity will be required, our findings suggest that the capacity of the Sphagnum community to gain carbon remained stable despite hydrological changes.

Paleoecological and historical data as an important tool in ecosystem management.

In recent decades, it has been observed that most forest fires in Europe were caused by people. Extreme droughts, which are more often prolonged, can increase the risk of forest fires, not only in southern Europe but also, in Central Europe. Nonetheless, catastrophic fire events are not well recognized in the Central European Lowlands (CEL), where large forest complexes are located. Knowledge of past fire activity in this part of Europe is scarce, although several fires have occurred in this area during the previous millennia. Large coniferous forest monocultures located in the CEL are highly susceptible to fires and other disturbances. Here, we present a case study from the Tuchola Pinewoods (TP; northern Poland), where large pine monocultures are present. The main aim of this study is to document the potential effects past land management has on modern day disturbance regimes using state-of-the-art paleoecological data, historical documents and cartographic materials. We then present a protocol that will help forest managers utilize long-term paleoecological records. Based on paleoecological investigations, historical documents, and cartographic materials, our results show that, in the past 300 years, the TP witnessed not only disastrous fires and but also windfalls by tornados and insect outbreaks. A change in management from Polish to Prussian/German in the 18th century led to the transformation of mixed forests into Scots pine monocultures with the purpose to allow better economic use of the forest. Those administrative decisions led to an ecosystem highly susceptible to disturbances. This article provides a critical review of past forest management as well as future research directions related to the impacts of fire risk on land management and ecosystem services: (a) habitat composition and structure (biodiversity); (b) natural water management; and (c) mitigation of climate changes. Designated forest conditions, management, and future fire risk are a controversial and highly debated topic of forest management by Forestry Units. More research will allow the gathering of reliable information pertinent to management practices with regard to the current fire risks. It is necessary to develop a dialog between scientists and managers to reduce the risk of fires in projected climate change.

Genetic Determinism vs. Phenotypic Plasticity in Protist Morphology.

Untangling the relationships between morphology and phylogeny is key to building a reliable taxonomy, but is especially challenging for protists, where the existence of cryptic or pseudocryptic species makes finding relevant discriminant traits difficult. Here we use Hyalosphenia papilio (a testate amoeba) as a model species to investigate the contribution of phylogeny and phenotypic plasticity in its morphology. We study the response of H. papilio morphology (shape and pores number) to environmental variables in (i) a manipulative experiment with controlled conditions (water level), (ii) an observational study of a within-site natural ecological gradient (water level), and (iii) an observational study across 37 European peatlands (climate). We showed that H. papilio morphology is correlated to environmental conditions (climate and water depth) as well as geography, while no relationship between morphology and phylogeny was brought to light. The relative contribution of genetic inheritance and phenotypic plasticity in shaping morphology varies depending on the taxonomic group and the trait under consideration. Thus, our data call for a reassessment of taxonomy based on morphology alone. This clearly calls for a substantial increase in taxonomic research on these globally still under-studied organisms leading to a reassessment of estimates of global microbial eukaryotic diversity.

Response of sphagnum peatland testate amoebae to a 1-year transplantation experiment along an artificial hydrological gradient.

Peatland testate amoebae (TA) are well-established bioindicators for depth to water table (DWT), but effects of hydrological changes on TA communities have never been tested experimentally. We tested this in a field experiment by placing Sphagnum carpets (15 cm diameter) collected in hummock, lawn and pool microsites (origin) at three local conditions (dry, moist and wet) using trenches dug in a peatland. One series of samples was seeded with microorganism extract from all microsites. TA community were analysed at T0: 8-2008, T1: 5-2009 and T2: 8-2009. We analysed the data using conditional inference trees, principal response curves (PRC) and DWT inferred from TA communities using a transfer function used for paleoecological reconstruction. Density declined from T0 to T1 and then increased sharply by T2. Species richness, Simpson diversity and Simpson evenness were lower at T2 than at T0 and T1. Seeded communities had higher species richness in pool samples at T0. Pool samples tended to have higher density, lower species richness, Simpson diversity and Simpson Evenness than hummock and/or lawn samples until T1. In the PRC, the effect of origin was significant at T0 and T1, but the effect faded away by T2. Seeding effect was strongest at T1 and lowest vanished by T2. Local condition effect was strong but not in line with the wetness gradient at T1 but started to reflect it by T2. Likewise, TA-inferred DWT started to match the experimental conditions by T2, but more so in hummock and lawn samples than in pool samples. This study confirmed that TA responds to hydrological changes over a 1-year period. However, sensitivity of TA to hydrological fluctuations, and thus the accuracy of inferred DWT changes, was habitat specific, pool TA communities being least responsive to environmental changes. Lawns and hummocks may be thus better suited than pools for paleoecological reconstructions. This, however, contrasts with the higher prediction error and species' tolerance for DWT with increasing dryness observed in transfer function models.

Neodymium isotopes in peat reveal past local environmental disturbances.

Over the past decade, the neodymium (Nd) isotope composition of mineral matter from peat cores has seen increasingly common use as a tracer of dust influx associated with major changes in the Holocene atmospheric circulation. However, the incomplete understanding of the local controls on the sources of the sediment supplied to peatlands remains a key difficulty in the interpretation of the archived Nd isotope signals. Here, we used neodymium isotopes to reconstruct environmental disturbances in peatlands. We performed a multi-proxy study of two peatlands that experienced peatland burning and validated the recorded peat Nd signatures using reference surface sampling. Our data show a link between the Nd isotope signals and local environmental disturbances: peat burning, local fire activity and pollution fluxes. Our study illustrates the crucial role of identifying local events that influence the supply of mineral material to peatlands. Insufficient recognition of such local controls may either obscure the large-scale variations in the atmospheric circulation patterns, or introduce artefacts to the Holocene climate record. We also provide recommendations for the use of Nd isotopes in palaeoecological studies of peatlands.

Synthesis of palaeoecological data from the Polish Lowlands suggests heterogeneous patterns of old-growth forest loss after the Migration Period.

Human impact on Central European forests dates back thousands of years. In this study we reanalyzed 36 published pollen data sets with robust chronologies from Polish Lowlands to determine the patterns of large-scale forest decline after the Migration Period (fourth to sixth century CE). The study revealed substantial heterogeneity in the old-growth forest decline patterns. Using new high-resolution studies, we could better understand the timing of this transition related to increasing economic development. After the Migration Period, forest expansion continued until the seventh to ninth centuries cal. CE, when the dawn of Slavic culture resulted in large-scale forest decline, especially in north-western and north-central Poland. Later, forest decline was recorded mainly in north-eastern Poland and was related to Prussian settlements, including activities associated with the Teutonic Order, as well as with new settlements from the fourteenth century. The composite picture shows a varied spatio-temporal forest loss and transition towards the present-day, human activity dominated landscapes. However, some sites, such as in north-eastern Poland, are characterized by a less abrupt critical transition. The pristine nature of the oak-hornbeam forest had already been destroyed in Early Medieval times (eighth to ninth centuries cal. CE) and the potential for recovery was largely lost. Our study has confirmed previous assumptions that the decline of hornbeam across the Polish Lowlands may be an early indicator of local settlement processes, preceding severe forest loss, and establishment of permanent agriculture.

Recent climate change has driven divergent hydrological shifts in high-latitude peatlands.

High-latitude peatlands are changing rapidly in response to climate change, including permafrost thaw. Here, we reconstruct hydrological conditions since the seventeenth century using testate amoeba data from 103 high-latitude peat archives. We show that 54% of the peatlands have been drying and 32% have been wetting over this period, illustrating the complex ecohydrological dynamics of high latitude peatlands and their highly uncertain responses to a warming climate.

How Joannites' economy eradicated primeval forest and created anthroecosystems in medieval Central Europe.

During European states' development, various past societies utilized natural resources, but their impact was not uniformly spatially and temporally distributed. Considerable changes resulted in landscape fragmentation, especially during the Middle Ages. Changes in state advances that affected the local economy significantly drove trajectories of ecosystems' development. The legacy of major changes from pristine forest to farming is visible in natural archives as novel ecosystems. Here, we present a high-resolution densely dated multi-proxy study covering the last 1500 years from a peatland located in CE Europe. The economic activity of medieval societies was highly modified by new rulers-the Joannites (the Order of St. John of Jerusalem, Knights Hospitaller). We studied the record of these directorial changes noted in the peat profile. Our research revealed a rapid critical land-use transition in the late Middle Ages and its consequences on the peatland ecosystem. The shift from the virgin forest with regular local fires to agriculture correlates well with the raising of local economy and deforestations. Along with the emerging openness, the wetland switched from alkaline wet fen state to acidic, drier Sphagnum-dominated peatland. Our data show how closely the ecological state of wetlands relates to forest microclimate. We identified a significant impact of the Joannites who used the novel farming organization. Our results revealed the surprisingly fast rate of how feudal economy eliminated pristine nature from the studied area and created novel anthroecosystems.

A novel testate amoebae trait-based approach to infer environmental disturbance in Sphagnum peatlands.

Species' functional traits are closely related to ecosystem processes through evolutionary adaptation, and are thus directly connected to environmental changes. Species' traits are not commonly used in palaeoecology, even though they offer powerful advantages in understanding the impact of environmental disturbances in a mechanistic way over time. Here we show that functional traits of testate amoebae (TA), a common group of palaeoecological indicators, can serve as an early warning signal of ecosystem disturbance and help determine thresholds of ecosystem resilience to disturbances in peatlands. We analysed TA traits from two Sphagnum-dominated mires, which had experienced different kinds of disturbances in the past 2000 years - fire and peat extraction, respectively. We tested the effect of disturbances on the linkages between TA community structure, functional trait composition and functional diversity using structural equation modelling. We found that traits such as mixotrophy and small hidden apertures (plagiostomic apertures) are strongly connected with disturbance, suggesting that these two traits can be used as palaeoecological proxies of peatland disturbance. We show that TA functional traits may serve as a good proxy of past environmental changes, and further analysis of trait-ecosystem relationships could make them valuable indicators of the contemporary ecosystem state.

Seasonal changes in Sphagnum peatland testate amoeba communities along a hydrological gradient.

Testate amoebae are an abundant and functionally important group of protists in peatlands, but little is known about the seasonal patterns of their communities. We investigated the relationships between testate amoeba diversity and community structure and water table depth and light conditions (shading vs. insolation) in a Sphagnum peatland in Northern Poland (Linje mire) in spring and summer 2010. We monitored the water table at five sites across the peatland and collected Sphagnum samples in lawn and hummock micro-sites around each piezometer, in spring (3 May) and mid-summer (6 August) 2010. Water table differed significantly between micro-sites and seasons (Kruskal-Wallis test, p=0.001). The community structure of testate amoebae differed significantly between spring and summer in both hummock and lawn micro-sites. We recorded a small, but significant drop in Shannon diversity, between spring and summer (1.76 vs. 1.72). Strongest correlations were found between testate amoeba communities and water table lowering and light conditions. The relative abundance of mixotrophic species Hyalosphenia papilio, Archerella flavum and of Euglypha ciliata was higher in the summer.