Vegetation and environmental changes in Northern Anatolia between 134 and 119 ka recorded in Black Sea sediments

This multiproxy study on SE Black Sea sediments provides the first detailed reconstruction of vegetation and environmental history of Northern Anatolia between 134 and 119 ka. Here, the glacial–interglacial transition is characterized by several short-lived alternating cold and warm events preceding a meltwater pulse (~ 130.4–131.7 ka). The latter is reconstructed as a cold arid period correlated to Heinrich event 11. The initial warming is evidenced at ~ 130.4 ka by increased primary productivity in the Black Sea, disappearance of ice-rafted detritus, and spreading of oaks in Anatolia. A Younger Dryas-type event is not identifiable. The Eemian vegetation succession corresponds to the main climatic phases in Europe: i) the Quercus–Juniperus phase (128.7–126.4 ka) indicates a dry continental climate; ii) the Ostrya–Corylus–Quercus–Carpinus phase (126.4–122.9 ka) suggests warm summers, mild winters, and high year-round precipitation; iii) the Fagus–Carpinus phase (122.9–119.5 ka) indicates cooling and high precipitation; and iv) increasing Pinus at ~ 121 ka marks the onset of cooler/drier conditions. Generally, pollen reconstructions suggest altitudinal/latitudinal migrations of vegetation belts in Northern Anatolia during the Eemian caused by increased transport of moisture. The evidence for the wide distribution of Fagus around the Black Sea contrasts with the European records and is likely related to climatic and genetic factors.

Climatic drivers of hourly to yearly tree radius variations along a 6°C natural warming gradient

Climate affects the timing, rate and dynamics of tree growth, over time scales ranging from seconds to centuries. Monitoring how a tree's stem radius varies over these time scales can provide insight into intra-annual stem dynamics and improve our understanding of climate impacts on tree physiology and growth processes. Here, we quantify the response of radial conifer stem size to environmental fluctuations via a novel assessment of tree circadian cycles. We analyze four years of sub-hourly data collected from 56 larch and spruce trees growing along a natural temperature gradient of ∼6 °C in the central Swiss Alps. During the growing season, tree stem diameters were greatest at mid-morning and smallest in the late evening, reflecting the daily cycle of water uptake and loss. Along the gradient, amplitudes calculated from the stem radius cycle were ∼50% smaller at the upper site (∼2200 m a.s.l.) relative to the lower site (∼800 m a.s.l.). We show changes in precipitation, temperature and cloud cover have a substantial effect on typical growing season diurnal cycles; amplitudes were nine times smaller on rainy days (>10 mm), and daily amplitudes are approximately 40% larger when the mean daily temperature is 15–20 °C than when it is 5–10 °C. We find that over the growing season in the sub-alpine forests, spruce show greater daily stem water movement than larch. However, under projected future warming, larch could experience up to 50% greater stem water use, which may severely affect future growth on already dry sites. Our data further indicate that because of the confounding influences of radial growth and short-term water dynamics on stem size, conventional methodology probably overstates the effect of water-linked meteorological variables (i.e. precipitation and relative humidity) on intra-annual tree growth. We suggest future studies use intra-seasonal measurements of cell development and consider whether climatic factors produce reversible changes in stem diameter. These study design elements may help researchers more accurately quantify and attribute changes in forest productivity in response to future warming.

The effect of abrupt climatic warming on biogeochemical cycling and N2O emissions in a terrestrial ecosystem

The large, rapid increase in atmospheric N2O concentrations that occurred concurrent with the abrupt warming at the end of the Last Glacial period might have been the result of a reorganization in global biogeochemical cycles. To explore the sensitivity of nitrogen cycling in terrestrial ecosystems to abrupt warming, we combined a scenario of climate and vegetation composition change based on multiproxy data for the Oldest Dryas–Bølling abrupt warming event at Gerzensee, Switzerland, with a biogeochemical model that simulates terrestrial N uptake and release, including N2O emissions. As for many central European sites, the pollen record at the Gerzensee is remarkable for the abundant presence of the symbiotic nitrogen fixer Hippophaë rhamnoides (L.) during the abrupt warming that also marks the beginning of primary succession on immature glacial soils. Here we show that without additional nitrogen fixation, climate change results in a significant increase of N2O emissions of approximately factor 3.4 (from 6.4 ± 1.9 to 21.6 ± 5.9 mg N2O–N m− 2 yr− 1). Each additional 1000 mg m− 2 yr− 1 of nitrogen added to the ecosystem through N-fixation results in additional N2O emissions of 1.6 mg N2O–N m− 2 yr− 1 for the time with maximum H. rhamnoides coverage. Our results suggest that local reactions of emissions to abrupt climate change could have been considerably faster than the overall atmospheric concentration changes observed in polar ice. Nitrogen enrichment of soils due to the presence of symbiotic N-fixers during early primary succession not only facilitates the establishment of vegetation on soils in their initial stage of development, but can also have considerable influence on biogeochemical cycles and the release of reactive nitrogen trace gases to the atmosphere.

Vegetation responses to rapid warming and to minor climatic fluctuations during the Late-Glacial Interstadial (GI-1) at Gerzensee (Switzerland)

High-resolution pollen analyses made on the same samples on which the ratios of oxygen isotopes were measured that provided the time scale and a temperature proxy after correlation to NorthGRIP. (1) A primary succession: The vegetation responded to the rapid rise of temperatures around 14,685 yr BP, with a primary succession on a decadal to centennial time scale. The succession between ca 15,600 and 13,000 yr BP included: (1.1.) The replacement of shrub-tundra by woodland of Juniperus and tree birch (around 14,665 yr BP) (1.2.) The response of Juniperus pollen to the shift in oxygen isotopes in less than 20 yr, (1.3.) A sequence of population increases of Hippophaë rhamnoides (ca 14,600 yr BP), Salix spp. (ca 14,600 yr BP), Betula trees (ca.14,480 yr BP), Populus cf. tremula (ca. 14,300 yr BP), and Pinus cf. sylvestris (ca. 13,830 yr BP). (2) Biological processes: Plants responded to the rapid increase of summer temperatures on all organisational levels: (2.1) Individuals may have produced more pollen (e.g. Juniperus); (2.2) Populations increased or decreased (e.g. Juniperus, Betula, later Pinus), and (2.3) Populations changed their biogeographical range and may show migrational lags. (2.4) Plant communities changed in their composition because the species pools changed through immigration and (local) extinction. Some plant communities may have been without modern analogue.These mechanisms require increasing amounts of time. (2.5) Processes on the level of ecosystems, with species interactions, may involve various time scales. Besides competition and facilitation, nitrogen fixation is discussed. (3) The minor fluctuations of temperature during the Late-Glacial Interstadial, which are recorded in δ18O, resulted in only very minor changes in pollen during the Aegelsee Oscillation (Older Dryas biozone, GI-1d) and the Gerzensee Oscillation (GI-1b). (4) Biodiversity: The afforestation at the onset of Bølling coincided with a gradual increase of taxonomic diversity up to the time of the major Pinus expansion.

A model-data comparison of Holocene timberline changes in the Swiss Alps reveals past and future drivers of mountain forest dynamics

Mountain vegetation is strongly affected by temperature and is expected to shift upwards with climate change. Dynamic vegetation models are often used to assess the impact of climate on vegetation and model output can be compared with paleobotanical data as a reality check. Recent paleoecological studies have revealed regional variation in the upward shift of timberlines in the Northern and Central European Alps in response to rapid warming at the Younger Dryas/Preboreal transition ca. 11700years ago, probably caused by a climatic gradient across the Alps. This contrasts with previous studies that successfully simulated the early Holocene afforestation in the (warmer) Central Alps with a chironomid-inferred temperature reconstruction from the (colder) Northern Alps. We use LandClim, a dynamic landscape vegetation model to simulate mountain forests under different temperature, soil and precipitation scenarios around Iffigsee (2065m a.s.l.) a lake in the Northwestern Swiss Alps, and compare the model output with the paleobotanical records. The model clearly overestimates the upward shift of timberline in a climate scenario that applies chironomid-inferred July-temperature anomalies to all months. However, forest establishment at 9800 cal. BP at Iffigsee is successfully simulated with lower moisture availability and monthly temperatures corrected for stronger seasonality during the early Holocene. The model-data comparison reveals a contraction in the realized niche of Abies alba due to the prominent role of anthropogenic disturbance after ca. 5000 cal. BP, which has important implications for species distribution models (SDMs) that rely on equilibrium with climate and niche stability. Under future climate projections, LandClim indicates a rapid upward shift of mountain vegetation belts by ca. 500m and treeline positions of ca. 2500m a.s.l. by the end of this century. Resulting biodiversity losses in the alpine vegetation belt might be mitigated with low-impact pastoralism to preserve species-rich alpine meadows.

A compilation of Western European terrestrial records 60–8 ka BP: towards an understanding of latitudinal climatic gradients

Terrestrial records of past climatic conditions, such as lake sediments and speleothems, provide data of great importance for understanding environmental changes. However, unlike marine and ice core records, terrestrial palaeodata are often not available in databases or in a format that is easily accessible to the non-specialist. As a consequence, many excellent terrestrial records are unknown to the broader palaeoclimate community and are not included in compilations, comparisons, or modelling exercises. Here we present a compilation of Western European terrestrial palaeo-records covering, entirely or partially, the 60–8-ka INTIMATE time period. The compilation contains 56 natural archives, including lake records, speleothems, ice cores, and terrestrial proxies in marine records. The compilation is limited to include records of high temporal resolution and/or records that provide climate proxies or quantitative reconstructions of environmental parameters, such as temperature or precipitation, and that are of relevance and interest to a broader community. We briefly review the different types of terrestrial archives, their respective proxies, their interpretation and their application for palaeoclimatic reconstructions. We also discuss the importance of independent chronologies and the issue of record synchronization. The aim of this exercise is to provide the wider palaeo-community with a consistent compilation of high-quality terrestrial records, to facilitate model-data comparisons, and to identify key areas of interest for future investigations. We use the compilation to investigate Western European latitudinal climate gradients during the deglacial period and, despite of poorly constrained chronologies for the older records, we summarize the main results obtained from NW and SW European terrestrial records before the LGM.

Human and climatic impact on mires: a case study of Les Amburnex mire, Swiss Jura Mountains

Modern period long-term human and climatic impacts on a small mire in the Jura Mountains were assessed using testate amoebae, macrofossils and pollen. This multiproxy data analysis permitted detailed interpretations of local and regional environmental change and thus a partial disentanglement of the different variables that influence long-term mire development. From the Middle Ages until a.d. 1700 the mire vegetation was characterised by ferns, Caltha and Vaccinium, but then abruptly changed into the modern vegetation characterised by Cyperaceae, Potentilla and Sphagnum. The cause for this change was most probably deforestation, possibly enhanced by climatic cooling. A decrease in trampling intensity by domestic animals from a.d. 1950 onwards allowed Sphagnum growth and climatic warming in the a.d. 1980s and 1990s may have been responsible for considerable changes in the species composition. The mire investigated is an example of the rapid changes in mire vegetation and peat development that occurred throughout the central European mountain region during the past centuries as a result of changing climate and land-use practice. These processes are still active today and will determine the future development of high-altitude mires.

Rapid responses of high-mountain vegetation to early Holocene environmental changes in the Swiss Alps

1 The Early Holocene sediment of a lake at tree line (Gouillé Rion, 2343 m a.s.l.) in the Swiss Central Alps was sampled for plant macrofossils. Thin (0.5 cm) slices, representing time intervals of c. 50 years each from 11 800 to 7800 cal. year bp, were analysed and the data compared with independent palaeoclimatic proxies to study vegetational responses to environmental change. 2 Alpine plant communities (e.g. with Salix herbacea) were established at 11 600–11 500 cal. year bp, when oxygen-isotope records showed that temperatures increased by c. 3–4 °C within decades. Larix decidua trees reached the site at c. 11 350 cal. year bp, probably in response to further warming by 1–2 °C. Forests dominated by L. decidua persisted until 9600 cal. year bp, when Pinus cembra became more important. 3 The dominance of Larix decidua for two millennia is explained by dry summer conditions, and possibly low winter temperatures, which favoured it over the late-successional Pinus cembra. Environmental conditions were a result of variations in the earth's orbit, leading to a maximum of summer and a minimum of winter solar radiation. Other heliophilous and drought-adapted species, such as Dryas octopetala and Juniperus nana, could persist in the open L. decidua forests, but were out-competed when the shade-tolerant P. cembra expanded. 4 The relative importance of Larix decidua decreased during periods of diminished solar radiation at 11 100, 10 100 and 9400 cal. year bp. Stable concentrations of L. decidua indicate that these percentage oscillations were caused by temporary increases of Pinus cembra, Dryas octopetala and Juniperus nana that can be explained by increases in moisture and/or decreases in summer temperature. 5 The final collapse of Larix decidua at 8400 cal. year bp was possibly related to abrupt climatic cooling as a consequence of a large meltwater input to the North Atlantic. Similarly, the temporary exclusion of Pinus cembra from tree line at 10 600–10 200 cal. year bp may be related to slowing down of thermohaline circulation at 10 700–10 300 cal. year bp. 6 Our results show that tree line vegetation was in dynamic equilibrium with climate, even during periods of extraordinarily rapid climatic change. They also imply that forecasted global warming may trigger rapid upslope movements of the tree line of up to 800 m within a few decades or centuries at most, probably inducing large-scale displacements of plant species as well as irrecoverable biodiversity losses.

Plant regeneration directs changes in grassland composition after extreme drought: a 13-year study in southern Switzerland

1. The cover of plant species was recorded annually from 1988 to 2000 in nine spatially replicated plots in a species-rich, semi-natural meadow at Negrentino (southern Alps). This period showed large climatic variation and included the centennial maximum and minimum frequency of days with ≥ 10 mm of rain. 2. Changes in species composition were compared between three 4-year intervals characterized by increasingly dry weather (1988–91), a preceding extreme drought (1992–95), and increasingly wet weather (1997–2000). Redundancy analysis and anova with repeated spatial replicates were used to find trends in vegetation data across time. 3. Recruitment capacity, the potential for fast clonal growth and seasonal expansion rate were determined for abundant taxa and tested in general linear models (GLM) as predictors for rates of change in relative cover of species across the climatically defined 4-year intervals. 4. Relative cover of the major growth forms present, graminoids and forbs, changed more in the period following extreme drought than at other times. Recruitment capacity was the only predictor of species’ rates of change. 5. Following perturbation, re-colonization was the primary driver of vegetation dynamics. The dominant grasses, which lacked high recruitment from seed, therefore decreased in relative abundance. This effect persisted until the end of the study and may represent a lasting response to an extreme climatic event.

The Early Spörer Minimum – A Period of Extraordinary Climate and Socio-economic Changes in Western and Central Europe

Throughout the last millennium, mankind was affected by prolonged deviations from the climate mean state. While periods like the Maunder Minimum in the 17th century have been assessed in greater detail, earlier cold periods such as the 15th century received much less attention due to the sparse information available. Based on new evidence from different sources ranging from proxy archives to model simulations, it is now possible to provide an end-to-end assessment about the climate state during an exceptionally cold period in the 15th century, the role of internal, unforced climate variability and external forcing in shaping these extreme climatic conditions, and the impacts on and responses of the medieval society in Central Europe. Climate reconstructions from a multitude of natural and human archives indicate that, during winter, the period of the early Spörer Minimum (1431–1440 CE) was the coldest decade in Central Europe in the 15th century. The particularly cold winters and normal but wet summers resulted in a strong seasonal cycle that challenged food production and led to increasing food prices, a subsistence crisis, and a famine in parts of Europe. As a consequence, authorities implemented adaptation measures, such as the installation of grain storage capacities, in order to be prepared for future events. The 15th century is characterised by a grand solar minimum and enhanced volcanic activity, which both imply a reduction of seasonality. Climate model simulations show that periods with cold winters and strong seasonality are associated with internal climate variability rather than external forcing. Accordingly, it is hypothesised that the reconstructed extreme climatic conditions during this decade occurred by chance and in relation to the partly chaotic, internal variability within the climate system.

Long-term Responses of Mountain Ecosystems to Environmental Changes: Resilience, Adjustment, and Vulnerability

The steep environmental gradients of mountain ecosystems over short distances reflect large gradients of several climatic parameters and hence provide excellent possibilities for ecological research on the effects of environmental change. To gain a better understanding of the dynamics of abiotic and biotic parameters of mountain ecosystems, long-term records are required since permanent plots in mountain regions cover in the best case about 50 - 70 years. In order to extend investigations of ecological dynamics beyond these temporal limitations of permanent plots, paleoecological approaches can be used if the sampling resolution can be adapted to ecological research questions, e.g. a sample every 10 years. Paleoecological studies in mountain ecosystems can provide new ecological insights through the combination of different spatial and temporal scales. [f we thus improve our understanding of processes across both steep environmental gradients and different time scales, we may be able to better estimate ecosystem responses to current and future environmental change (Ammann et al. 1993; Lotter et al. 1997). The complexity of ecological interactions in mountain regions forces us to concentrate on a number of sub-systems - without losing sight of the wider context. Here, we summarize a few case studies on the effects of Holocene climate change and disturbance on the vegetation of the Western Alps. To categorize the main response modes of vegetation to climatic change and disturbance in the Alps we use three classes of ecological behaviour: "resilience", "adjustment", and "vulnerability", We assume a resilient (or elastic) behaviour if vegetation is able to recover to its former state, regaining important ecosystem characteristics, such as floristic composition, biodiversity, species abundances, and biomass (e.g. Küttel 1990; Aber and Melillo 199 1). Conversely, vegetation displacements may occur in response to climatic change and/or disturbance. In some cases, this may culminate in irreversible large-scale processes such as species and/or community extinctions. Such drastic developments indicate high ecosystem vulnerability (or inelasticity or instability, for detailed definitions see Küttel 1990; Aber and Melillo 199 1) to climatic change and/or disturbance. In this sense, the "vulnerability" (or instability) of an ecosystem is expressed by the degree of failure to recover to the original state before disturbance and/or climatic change. Between these two extremes (resilience vs. vulnerability), ecosystem adjustments to climatic change and/or disturbance may occur, including the appearance of new and/or the disappearance of old species. The term "adjustment" is hence used to indicate the response of vegetational communities, which adapted to new environmental conditions without losing their main character. For forest ecosystems, we assume vegetational adjustments (rather than vulnerability) if the dominant (or co-dominant) tree species are not outnumbered or replaced by formerly unimportant plant species or new invaders. Adaptation as a genetic process is not discussed here and will require additional pbylogeographical studies (that incorporate the analysis of ancient DNA) in order to fully understand the distributions of ecotypes.