No abstract

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.

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.

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.