Dr. Anita Nussbaumer

Many perennial plants show mast seeding, characterized by synchronous and highly variable reproduction across years. We propose a general model of masting, integrating proximate factors (environmental variation, weather cues, and resource budgets) with ultimate drivers (predator satiation and pollination efficiency). This general model shows how the relationships between masting and weather shape the diverse responses of species to climate warming, ranging from no change to lower interannual variation or reproductive failure. The role of environmental prediction as a masting driver is being reassessed; future studies need to estimate prediction accuracy and the benefits acquired. Since reproduction is central to plant adaptation to climate change, understanding how masting adapts to shifting environmental conditions is now a central question.

Climate change effects on tree reproduction are poorly understood, even though the resilience of populations relies on sufficient regeneration to balance increasing rates of mortality. Forest-forming tree species often mast, i.e. reproduce through synchronised year-to-year variation in seed production, which improves pollination and reduces seed predation. Recent observations in European beech show, however, that current climate change can dampen interannual variation and synchrony of seed production and that this masting breakdown drastically reduces the viability of seed crops. Importantly, it is unclear under which conditions masting breakdown occurs and how widespread breakdown is in this pan-European species. Here, we analysed 50 long-term datasets of population-level seed production, sampled across the distribution of European beech, and identified increasing summer temperatures as the general driver of masting breakdown. Specifically, increases in site-specific mean maximum temperatures during June and July were observed across most of the species range, while the interannual variability of population-level seed production (CVp) decreased. The declines in CVp were greatest, where temperatures increased most rapidly. Additionally, the occurrence of crop failures and low seed years has decreased during the last four decades, signalling altered starvation effects of masting on seed predators. Notably, CVp did not vary among sites according to site mean summer temperature. Instead, masting breakdown occurs in response to warming local temperatures (i.e. increasing relative temperatures), such that the risk is not restricted to populations growing in warm average conditions. As lowered CVp can reduce viable seed production despite the overall increase in seed count, our results warn that a covert mechanism is underway that may hinder the regeneration potential of European beech under climate change, with great potential to alter forest functioning and community dynamics.

Habitat maps at high thematic and spatial resolution and broad extents are fundamental tools for biodiversity conservation, the planning of ecological networks and the management of ecosystem services. To derive a habitat map for Switzerland, we used a composite methodology bringing together the best available spatial data and distribution models. The approach relies on the segmentation and classification of high spatial resolution (1 m) aerial imagery. Land cover data, as well as habitat and species distribution models built on Earth observation data from Sentinel 1 and 2, Landsat, Planetscope and LiDAR, inform the rule-based classification to habitats defined by the hierarchical Swiss Habitat Typology (TypoCH). A total of 84 habitats in 32 groups and 9 overarching classes are mapped in a spatially explicit manner across Switzerland. Validation and plausibility analysis with four independent datasets show that the mapping is broadly plausible, with good accuracy for most habitats, although with lower performance for fine-scale and linear habitats, habitats with restricted geographical distributions and those predominantly characterised by understorey species, especially forest habitats. The resulting map is a vector dataset available for interactive viewing and download from open EnviDat data sharing platform. The methodology is semi-automated to allow for updates over time.

The effects of tree pollen on precipitation chemistry are not fully understood and this can lead to misinterpretations of element deposition in European forests. We investigated the relationship between forest throughfall (TF) element fluxes and the Seasonal Pollen Integral (SPIn) using linear mixed-effects modelling (LME). TF was measured in 1990-2018 during the main pollen season (MPS, arbitrary two months) in 61 managed, mostly pure, even-aged Fagus, Quercus, Pinus, and Picea stands which are part of the ICP Forests Level II network. The SPIn for the dominant tree genus was observed at 56 aerobiological monitoring stations in nearby cities. The net contribution of pollen was estimated as the TF flux in the MPS minus the fluxes in the preceding and succeeding months. In stands of Fagus and Picea, two genera that do not form large amounts of flowers every year, TF fluxes of potassium (K⁺), ammonium-nitrogen (NH₄⁺-N), dissolved organic carbon (DOC), and dissolved organic nitrogen (DON) showed a positive relationship with SPIn. However- for Fagus- a negative relationship was found between TF nitrate-nitrogen (NO₃^--N) fluxes and SPIn. For Quercus and Pinus, two genera producing many flowers each year, SPIn displayed limited variability and no clear association with TF element fluxes. Overall, pollen contributed on average 4.1-10.6% of the annual TF fluxes of K⁺ > DOC > DON > NH₄⁺-N with the highest contribution in Quercus > Fagus > Pinus > Picea stands. Tree pollen appears to affect TF inorganic nitrogen fluxes both qualitatively and quantitatively, acting as a source of NH₄⁺-N and a sink of NO₃^--N. Pollen appears to play a more complex role in nutrient cycling than previously thought.

Bewirtschaftung und Umweltbedingungen beeinflussen die Gestalt von Wäldern. Mit der Einrichtung eines Naturwaldreservats wurden am Bettlachstock im Kanton Solothurn die Weichen zur Entwickung in Richtung eines Naturwaldes gestellt. Im Waldreservat werden im Rahmen des «Monitoring Naturwaldreservate Schweiz» mit Stichprobeninventuren die natürlichen Prozesse der Waldentwicklung und in der «Langfristigen Waldökosystemforschung (LWF)» mit kontinuierlichen Messungen der Einfluss von Luftverschmutzung und Klimawandel auf den Wald erforscht. Beides sind langfristige Untersuchungen, die Methoden der Waldinventur und Erfassung des Waldzustandes verwenden. Das Reservat ist aufgrund der Standortsvielfalt reich an verschiedenen Baumarten. Diese widerspiegeln das natürliche Artenspektrum. Der hohe Anteil an Fichten zeigt jedoch, dass die frühere Bewirtschaftung diese bewusst gefördert hat. Die Luftbelastung und das Klima hatten und haben nachweislich einen Einfluss auf den Wald am Bettlachstock. Die Die Luftbelastung ist eher rückläufig und die Klimaveränderung eher zunehmend. Beide stehen aus heutiger Sicht grundsätzlich der Entwicklung zu einem naturnahen Wald an diesem Standort nicht im Wege. Eine Zunahme von trockenen Sommern ist zu erwarten und könnte die Standorteigenschaften verändern. In Teilen des Reservats könnte dies mittelfristig zu vermehrtem Absterben von Tannen und Fichten und in der Folge zu einer Zunahme des Totholzvolumens führen. Langfristig könnte dies Verschiebungen der Höhenstufen und in der Baumartenzusammensetzung hin zu mehr Laubbaumarten auslösen.

Resource allocation to different plant tissues is likely to be affected by high investment into fruit production during mast years. However, there is a large knowledge gap concerning species-specific differences in resource dynamics. We investigated the influence of mast years on stem growth, leaf production, and leaf carbon (C), nitrogen (N), and phosphorus (P) concentrations and contents in Fagus sylvatica, Quercus petraea, and Q. robur at continental and climate region scales using long-term data from the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) and similar datasets. We discussed the results in the light of opposing resource dynamics hypotheses: (i) resource accumulation before mast years and exhaustion after mast years (resource storage hypothesis), (ii) shifting resources from vegetative to generative compartments (resource switching hypothesis), and (iii) investing resources concurrently in both vegetative and generative compartments (resource matching hypothesis). Linear mixed-effects modelling (LMM) showed that both stem growth and leaf production were negatively influenced by weather conditions which simultaneously lead to high fruit production. Thus, the impact of generative on vegetative growth is intermixed with effects of environmental factors. Superposed epoch analyses and LMM showed that for mast behaviour in F. sylvatica, there are indicators supporting the resource storage and the resource switching hypotheses. Before mast years, resources were accumulated, while during mast years resources switched from vegetative to generative tissues with reduced stem and leaf growth. For the Quercus species, stem growth was reduced after mast years, which supports the resource storage hypothesis. LMM showed that leaf C concentrations did not change with increasing fruit production in neither species. Leaf N and P concentrations increased in F. sylvatica, but not in Quercus species. Leaf N and P contents decreased with increasing fruit production in all species, as did leaf C content in F. sylvatica. Overall, our findings suggest different resource dynamics strategies in F. sylvatica and Quercus species, which might lead to differences in their adaptive capacity to a changing climate.

Die Rotbuche (Fagus sylvatica) zeigt mit ihrer Fortpflanzungsstrategie ein klares Mastverhalten, das heisst, sie produziert nur in einzelnen Jahren sehr grosse Mengen an Blüten und Früchten (Bucheckern). Bisher wurde angenommen, dass die Buche zu den Baumarten gehört, die bei erfolgter Bestäubung in aller Regel auch Früchte produziert. Im Schweizer Hitze- und Trockensommer 2018 reagierte die Buchen an einigen Standorten aber anders: Nach einer sehr starken Blüte entwickelten sie nur wenige Früchte, oder sie warfen die unvollständig entwickelten Früchte bereits Mitte des Sommers ab. Die Analyse von Messdaten der letzten 15 bis 19 Jahre von drei Buchenuntersuchungsflächen der Langfristigen Waldökosystem-Forschung (LWF) zeigt, dass starke Hitzewellen und langanhaltende Trockenheit während der Fruchtentwicklung zu einem Abbruch der Fruchtproduktion führen können. Diese extremen Wetterverhältnisse funktionieren somit als «Umweltveto». In Jahren mit vergleichbaren Bedingungen während der Fruchtentwicklung waren die Durchschnittstemperaturen im Sommer um 1.5 °C höher, und der Niederschlag war um 45% niedriger als im langjährigen Mittel. Der angenommene Zweijahreszyklus der Buchenblüte kann somit nicht nur durch ungünstige Wetterbedingungen während der Bestäubung, sondern auch während der Fruchtentwicklung unterbrochen werden.

Years with high fruit production, known as mast years, are the usual reproduction strategy of European beech. Harsh weather conditions such as frost during flowering can lead to pollination failure in spring. It has been assumed that mast is controlled by flowering, and that after successful pollination, high amounts of fruits and seeds would be produced. However, the extremely hot and dry European summer of 2018 showed that despite successful pollination, beechnuts did not develop or were only abundant in a few forest stands. An in-depth analysis of three forest sites of European beech from the Swiss Long-Term Forest Ecosystem Research Programme over the last 15-19 years revealed for the first time that extreme summer heat and drought can act as an "environmental veto", leading to early fruit abortion. Within the forest stands in years with fruit abortion, summer mean temperatures were 1.5 °C higher and precipitation sums were 45% lower than the long-term average. Extreme summer heat and drought, together with frost during flowering, are therefore disrupting events of the assumed biennial fruiting cycle in European beech.

Synchronisation of seed production is the reproductive strategy of many long-lived plant species. Years with high seed production, so called mast years, impact various ecological and economical areas. It e.g. provides additional nutrition for wildlife populations and livestock, can increase infection pressure of zoonoses, and affects timber production. Furthermore, mast years are likely to lead to changes in resource allocation to vegetative and generative tissues. For many species, mast years can be triggered by weather conditions in the years before and during the mast year. In the light of the current climate change, spatial and temporal mast patterns are, therefore, likely to change. Hence, improving the knowledge on this phenomenon is crucial for future forest ecosystem research, forest management and forest modelling. [...]

Die Synchronisierung der Samenproduktion ist für viele langlebige Pflanzenarten eine erfolgreiche Fortpflanzungsstrategie. Jahre mit starker Samenproduktion, so genannte Mastjahre, wirken sich auf verschiedene ökologische und ökonomische Bereiche aus. Mastjahre bieten z.B. zusätzliche Nahrung für Wildtierpopulationen und Nutztiere, können den Infektionsdruck von Zoonosen erhöhen und haben einen Einfluss auf die Holzproduktion. Darüber hinaus können Mastjahre zu Veränderungen der Ressourcenallokation zwischen vegetativen und generativen Geweben führen. Bei vielen Arten werden Mastjahre durch Wetterbedingungen in den Jahren vor und während des Mastjahres ausgelöst. Es kann davon ausgegangen werden, dass sich räumliche und zeitliche Mastmuster angesichts des aktuellen Klimawandels verändern werden. Daher ist ein besseres Verständnis dieses Phänomens von entscheidender Bedeutung für künftige Waldökosystemforschung, Waldbewirtschaftung sowie Waldmodellierung. [...]

Mast seeding, the synchronised occurrence of large amounts of fruits and seeds at irregular intervals, is a reproductive strategy in many wind-pollinated species. Although a series of studies have investigated mast year (MY) patterns in European forest tree species at the regional scale, there are few recent evaluations at a European scale on the impact of weather variables (weather cues) and resource dynamics on mast behaviour. Thus the main objective of this study is to investigate the impact of specific weather conditions, as environmental drivers for MYs, on resources in Fagus sylvatica L., Quercus petraea (MATT.) LIEBL., Quercus robur L., Picea abies (L.) KARST. and Pinus sylvestris L. at a European level and to explore the robustness of the relationships in smaller regions within Europe. Data on seed production originating from the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) were analysed. Three beta regression models were applied to investigate the impact of seasonal weather variables on MY occurrence, as well as the influence of fruiting intensity levels in the years prior to MYs. Resource dynamics are analysed at three different spatial scales (continent, countries and ecoregions).
At a European scale, important weather cues for beech MYs were a cold and wet summer two years before a MY, a dry and warm summer one year before a MY and a warm spring in the MY. For spruce, a cold and dry summer two years prior to a MY and a warm and dry summer in the year before the MY showed the strongest associations with the MY. For oak, high spring temperature in the MY was the most important weather cue. For beech and spruce, and to some extent also for oak species, the best fitting models at European scale were well reflected by those found at smaller scales. For pine, best fitting models were highly diverse concerning weather cues. Fruiting levels were high in all species two years before the MY and also high one year before the MY in the oak species and in pine. In beech, fruiting levels one year before the MY were not important and in spruce, they were inconsistent depending on the region. As a consequence, evidence of resource depletion could only be seen in some regions for spruce.

Occurrence of mast years, i.e. the synchronous production of vast amounts of fruits or seeds, has an important impact on forest ecosystems, their functioning and their services. We investigated the mast patterns of the forest tree species common beech, common and sessile oak, Norway spruce and Scots pine in Central and Northern Europe over the last two to three decades. We analysed data from the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) and additional Danish, German, Flemish and Swiss datasets.
Within-plot synchrony of fructification intensity in individual trees was high in beech and spruce and lower in oak species and pine. Mast frequency increased in most regions for beech, whereas the other species showed mixed or no trends. Beech, oak species and spruce showed strong mast year (MY) synchrony, but pine did not. MY synchrony between species was only significant in Bavaria, in Switzerland and between beech, oak species and spruce in Denmark. The deciduous species showed bimodal normal masting, while the conifers had switching normal masting. Oak species and the conifers supported the large seed and the accessory costs hypotheses, and beech and spruce supported the economy of scale, predator satiation and resource allocation hypotheses.

Why is the annual fruit production in forest tree species not constant, and which factors cause massive fruit production (seed mast year)? These and other related questions were already posed more than 100 years ago when tree breeding was economically beneficial. The questions have not been fully answered, yet. Rather, the same questions are studied again today as the climate is changing and the uncertainty about the continuation of forests at their current locations is growing. A 25 year long observation series on the variation of fruit production in Switzerland revealed a mean frequency of three years for mast seeding (full and medium mast) at low elevation on the Central Plateau in European beech (Fagus sylvatica), oak (Quercus petraea, Q. robur) and silver fir (Abies alba). In contrast, mast seed years of Norway spruce (Picea abies) occurred, on average, every sixth year. In 1992 and 2011, all four species synchronously showed mast seeding. The results are discussed in the light of different theories and new research findings. From the state of the current know ledge, we derive the need for long-term and fine-scale baseline data and present the new reporting and information webpage «mast web.ch». Here, volunteers can report observations on the fruit production of main tree species following a few simple criteria (citizen science approach). With this data, distribution maps on mass fructification levels will be made available and will serve for spatio-temporal fine-scale studies on mast seeding phenomena.