Fennoscandian tree-ring anatomy shows a warmer modern than medieval climate.

Earth system models and various climate proxy sources indicate global warming is unprecedented during at least the Common Era1. However, tree-ring proxies often estimate temperatures during the Medieval Climate Anomaly (950-1250 CE) that are similar to, or exceed, those recorded for the past century2,3, in contrast to simulation experiments at regional scales4. This not only calls into question the reliability of models and proxies but also contributes to uncertainty in future climate projections5. Here we show that the current climate of the Fennoscandian Peninsula is substantially warmer than that of the medieval period. This highlights the dominant role of anthropogenic forcing in climate warming even at the regional scale, thereby reconciling inconsistencies between reconstructions and model simulations. We used an annually resolved 1,170-year-long tree-ring record that relies exclusively on tracheid anatomical measurements from Pinus sylvestris trees, providing high-fidelity measurements of instrumental temperature variability during the warm season. We therefore call for the construction of more such millennia-long records to further improve our understanding and reduce uncertainties around historical and future climate change at inter-regional and eventually global scales.

Precise date for the Laacher See eruption synchronizes the Younger Dryas.

The Laacher See eruption (LSE) in Germany ranks among Europe's largest volcanic events of the Upper Pleistocene1,2. Although tephra deposits of the LSE represent an important isochron for the synchronization of proxy archives at the Late Glacial to Early Holocene transition3, uncertainty in the age of the eruption has prevailed4. Here we present dendrochronological and radiocarbon measurements of subfossil trees that were buried by pyroclastic deposits that firmly date the LSE to 13,006 ± 9 calibrated years before present (BP; taken as AD 1950), which is more than a century earlier than previously accepted. The revised age of the LSE necessarily shifts the chronology of European varved lakes5,6 relative to the Greenland ice core record, thereby dating the onset of the Younger Dryas to 12,807 ± 12 calibrated years BP, which is around 130 years earlier than thought. Our results synchronize the onset of the Younger Dryas across the North Atlantic-European sector, preclude a direct link between the LSE and Greenland Stadial-1 cooling7, and suggest a large-scale common mechanism of a weakened Atlantic Meridional Overturning Circulation under warming conditions8-10.

Return of the moth: rethinking the effect of climate on insect outbreaks.

The sudden interruption of recurring larch budmoth (LBM; Zeiraphera diniana or griseana Gn.) outbreaks across the European Alps after 1982 was surprising, because populations had regularly oscillated every 8-9 years for the past 1200 years or more. Although ecophysiological evidence was limited and underlying processes remained uncertain, climate change has been indicated as a possible driver of this disruption. An unexpected, recent return of LBM population peaks in 2017 and 2018 provides insight into this insect's climate sensitivity. Here, we combine meteorological and dendrochronological data to explore the influence of temperature variation and atmospheric circulation on cyclic LBM outbreaks since the early 1950s. Anomalous cold European winters, associated with a persistent negative phase of the North Atlantic Oscillation, coincide with four consecutive epidemics between 1953 and 1982, and any of three warming-induced mechanisms could explain the system's failure thereafter: (1) high egg mortality, (2) asynchrony between egg hatch and foliage growth, and (3) upward shifts of outbreak epicentres. In demonstrating that LBM populations continued to oscillate every 8-9 years at sub-outbreak levels, this study emphasizes the relevance of winter temperatures on trophic interactions between insects and their host trees, as well as the importance of separating natural from anthropogenic climate forcing on population behaviour.

Improved recovery of ancient DNA from subfossil wood - application to the world's oldest Late Glacial pine forest.

Ancient DNA from historical and subfossil wood has a great potential to provide new insights into the history of tree populations. However, its extraction and analysis have not become routine, mainly because contamination of the wood with modern plant material can complicate the verification of genetic information. Here, we used sapwood tissue from 22 subfossil pines that were growing c. 13 000 yr bp in Zurich, Switzerland. We developed and evaluated protocols to eliminate surface contamination, and we tested ancient DNA authenticity based on plastid DNA metabarcoding and the assessment of post-mortem DNA damage. A novel approach using laser irradiation coupled with bleaching and surface removal was most efficient in eliminating contaminating DNA. DNA metabarcoding confirmed which ancient DNA samples repeatedly amplified pine DNA and were free of exogenous plant taxa. Pine DNA sequences of these samples showed a high degree of cytosine to thymine mismatches, typical of post-mortem damage. Stringent decontamination of wood surfaces combined with DNA metabarcoding and assessment of post-mortem DNA damage allowed us to authenticate ancient DNA retrieved from the oldest Late Glacial pine forest. These techniques can be applied to any subfossil wood and are likely to improve the accessibility of relict wood for genome-scale ancient DNA studies.

European springtime temperature synchronises ibex horn growth across the eastern Swiss Alps.

Direct effects of climate change on animal physiology, and indirect impacts from disruption of seasonal synchrony and breakdown of trophic interactions are particularly severe in Arctic and Alpine ecosystems. Unravelling biotic from abiotic drivers, however, remains challenging because high-resolution animal population data are often limited in space and time. Here, we show that variation in annual horn growth (an indirect proxy for individual performance) of 8043 male Alpine ibex (Capra ibex) over the past four decades is well synchronised among eight disjunct colonies in the eastern Swiss Alps. Elevated March to May temperatures, causing premature melting of Alpine snowcover, earlier plant phenology and subsequent improvement of ibex food resources, fuelled annual horn growth. These results reveal dependency of local trophic interactions on large-scale climate dynamics, and provide evidence that declining herbivore performance is not a universal response to global warming even for high-altitude populations that are also harvested.

Tree-rings reveal two strong solar proton events in 7176 and 5259 BCE.

The Sun sporadically produces eruptive events leading to intense fluxes of solar energetic particles (SEPs) that dramatically disrupt the near-Earth radiation environment. Such events have been directly studied for the last decades but little is known about the occurrence and magnitude of rare, extreme SEP events. Presently, a few events that produced measurable signals in cosmogenic radionuclides such as 14C, 10Be and 36Cl have been found. Analyzing annual 14C concentrations in tree-rings from Switzerland, Germany, Ireland, Russia, and the USA we discovered two spikes in atmospheric 14C occurring in 7176 and 5259 BCE. The ~2% increases of atmospheric 14C recorded for both events exceed all previously known 14C peaks but after correction for the geomagnetic field, they are comparable to the largest event of this type discovered so far at 775 CE. These strong events serve as accurate time markers for the synchronization with floating tree-ring and ice core records and provide critical information on the previous occurrence of extreme solar events which may threaten modern infrastructure.

Axial changes in wood functional traits have limited net effects on stem biomass increment in European beech (Fagus sylvatica).

During the growing season, trees allocate photoassimilates to increase their aboveground woody biomass in the stem (ABIstem). This 'carbon allocation' to structural growth is a dynamic process influenced by internal and external (e.g., climatic) drivers. While radial variability in wood formation and its resulting structure have been intensively studied, their variability along tree stems and subsequent impacts on ABIstem remain poorly understood. We collected wood cores from mature trees within a fixed plot in a well-studied temperate Fagus sylvatica L. forest. For a subset of trees, we performed regular interval sampling along the stem to elucidate axial variability in ring width (RW) and wood density (ρ), and the resulting effects on tree- and plot-level ABIstem. Moreover, we measured wood anatomical traits to understand the anatomical basis of ρ and the coupling between changes in RW and ρ during drought. We found no significant axial variability in ρ because an increase in the vessel-to-fiber ratio with smaller RW compensated for vessel tapering towards the apex. By contrast, temporal variability in RW varied significantly along the stem axis, depending on the growing conditions. Drought caused a more severe growth decrease, and wetter summers caused a disproportionate growth increase at the stem base compared with the top. Discarding this axial variability resulted in a significant overestimation of tree-level ABIstem in wetter and cooler summers, but this bias was reduced to ~2% when scaling ABIstem to the plot level. These results suggest that F. sylvatica prioritizes structural carbon sinks close to the canopy when conditions are unfavorable. The different axial variability in RW and ρ thereby indicates some independence of the processes that drive volume growth and wood structure along the stem. This refines our knowledge of carbon allocation dynamics in temperate diffuse-porous species and contributes to reducing uncertainties in determining forest carbon fixation.

Tree physiological monitoring of the 2018 larch budmoth outbreak: preference for leaf recovery and carbon storage over stem wood formation in Larix decidua.

Insect defoliation impacts forest productivity worldwide, highlighting the relevance of plant-insect interactions. The larch budmoth (Zeiraphera griseana Hübner) is one of the most extensively studied defoliators, where numerous tree ring-based analyses on its host (Larix decidua Mill.) have aided in identifying outbreak dynamics over the past millennia. Yet, outbreaks have been widely absent after the early 1980s, and little is known about the in situ tree physiological responses and the allocation of carbon resources during and after defoliation. In summer 2018, we tracked an ongoing larch budmoth outbreak in a well-studied larch forest in the Swiss Alps. We performed biweekly monitoring on an affected and unaffected site using a unique combination of xylogenesis observations, measurements of non-structural carbohydrates, isotopic analysis of needle assimilates and ground-based and remote-sensed leaf trait observations. The budmoth induced a defoliation that lasted 40 days and could be detected by satellite observations. Soluble sugars significantly decreased in needles and stem phloem of the defoliated trees, while starch levels remained stable in the stem and root xylem compared to the control. Carbon and oxygen isotope ratios in needle assimilates indicated that neither photosynthetic assimilation rates nor stomatal conductance was different between sites before, during and after the outbreak. Defoliated trees ceased cell wall thickening 17 days earlier than unaffected trees, showing the earliest halt of ring formation recorded from 2007 untill 2013 and causing significant thinner cell walls, particularly in the latewood. No significant differences were found for cell enlargement rates and ring width. Our study revealed that an outbreak causes a downregulation of cell wall thickening first, while no starch is mobilized or leaf physiology is adjusted to compensate for the reduced carbon source due to defoliation. Our observations suggest that affected larch trees prioritize leaf recovery and carbon storage over wood biomass development.

1200 years of regular outbreaks in alpine insects.

The long-term history of Zeiraphera diniana Gn. (the larch budmoth, LBM) outbreaks was reconstructed from tree rings of host subalpine larch in the European Alps. This record was derived from 47513 maximum latewood density measurements, and highlights the impact of contemporary climate change on ecological disturbance regimes. With over 1000 generations represented, this is the longest annually resolved record of herbivore population dynamics, and our analysis demonstrates that remarkably regular LBM fluctuations persisted over the past 1173 years with population peaks averaging every 9.3 years. These regular abundance oscillations recurred until 1981, with the absence of peak events during recent decades. Comparison with an annually resolved, millennium-long temperature reconstruction representative for the European Alps (r=0.72, correlation with instrumental data) demonstrates that regular insect population cycles continued despite major climatic changes related to warming during medieval times and cooling during the Little Ice Age. The late twentieth century absence of LBM outbreaks, however, corresponds to a period of regional warmth that is exceptional with respect to the last 1000+ years, suggesting vulnerability of an otherwise stable ecological system in a warming environment.

New Insights into the Complex Relationship between Weight and Maturity of Burgundy Truffles (Tuber aestivum).

Despite an increasing demand for Burgundy truffles (Tuber aestivum), gaps remain in our understanding of the fungus' overall lifecycle and ecology. Here, we compile evidence from three independent surveys in Hungary and Switzerland. First, we measured the weight and maturity of 2,656 T. aestivum fruit bodies from a three-day harvest in August 2014 in a highly productive orchard in Hungary. All specimens ranging between 2 and 755 g were almost evenly distributed through five maturation classes. Then, we measured the weight and maturity of another 4,795 T. aestivum fruit bodies harvested on four occasions between June and October 2015 in the same truffière. Again, different maturation stages occurred at varying fruit body size and during the entire fruiting season. Finally, the predominantly unrelated weight and maturity of 81 T. aestivum fruit bodies from four fruiting seasons between 2010 and 2013 in Switzerland confirmed the Hungarian results. The spatiotemporal coexistence of 7,532 small-ripe and large-unripe T. aestivum, which accumulate to ~182 kg, differs from species-specific associations between the size and ripeness that have been reported for other mushrooms. Although size-independent truffle maturation stages may possibly relate to the perpetual belowground environment, the role of mycelial connectivity, soil property, microclimatology, as well as other abiotic factors and a combination thereof, is still unclear. Despite its massive sample size and proof of concept, this study, together with existing literature, suggests consideration of a wider ecological and biogeographical range, as well as the complex symbiotic fungus-host interaction, to further illuminate the hidden development of belowground truffle fruit bodies.

A technical perspective in modern tree-ring research--how to overcome dendroecological and wood anatomical challenges.

Dendroecological research uses information stored in tree rings to understand how single trees and even entire forest ecosystems responded to environmental changes and to finally reconstruct such changes. This is done by analyzing growth variations back in time and correlating various plant-specific parameters to (for example) temperature records. Integrating wood anatomical parameters in these analyses would strengthen reconstructions, even down to intra-annual resolution. We therefore present a protocol on how to sample, prepare, and analyze wooden specimen for common macroscopic analyses, but also for subsequent microscopic analyses. Furthermore we introduce a potential solution for analyzing digital images generated from common small and large specimens to support time-series analyses. The protocol presents the basic steps as they currently can be used. Beyond this, there is an ongoing need for the improvement of existing techniques, and development of new techniques, to record and quantify past and ongoing environmental processes. Traditional wood anatomical research needs to be expanded to include ecological information to this field of research. This would support dendro-scientists who intend to analyze new parameters and develop new methodologies to understand the short and long term effects of specific environmental factors on the anatomy of woody plants.

Causes and consequences of past and projected Scandinavian summer temperatures, 500-2100 AD.

Tree rings dominate millennium-long temperature reconstructions and many records originate from Scandinavia, an area for which the relative roles of external forcing and internal variation on climatic changes are, however, not yet fully understood. Here we compile 1,179 series of maximum latewood density measurements from 25 conifer sites in northern Scandinavia, establish a suite of 36 subset chronologies, and analyse their climate signal. A new reconstruction for the 1483-2006 period correlates at 0.80 with June-August temperatures back to 1860. Summer cooling during the early 17th century and peak warming in the 1930s translate into a decadal amplitude of 2.9°C, which agrees with existing Scandinavian tree-ring proxies. Climate model simulations reveal similar amounts of mid to low frequency variability, suggesting that internal ocean-atmosphere feedbacks likely influenced Scandinavian temperatures more than external forcing. Projected 21st century warming under the SRES A2 scenario would, however, exceed the reconstructed temperature envelope of the past 1,500 years.