Accurate estimates of seed production are central for understanding mast seeding mechanisms at tree and forest scales, and for designing sustainable management strategies. As trees are long-lived organisms, a long-term perspective is required to understand how reproduction acts during the life cycle of a tree. However, long-term series of seed production are challenging to obtain, as the available seed count procedures strictly rely on field methods, which are cost- and time-consuming, inherently limiting their widespread use at extensive spatial and temporal scales. In this study, we proposed a simple, rapid and flexible field method based on counting the seed in mobile ground quadrats (GQ), which was tested in beech forests. Quadrat measurements were first validated against reference measurements obtained from litter traps (LT) in three permanent plots. Results indicated that GQ provides robust and reliable estimates of seeds, which are not affected by seed predation occurring at the forest floor. Additional quadrat measurements were performed to evaluate the influence of sampling schemes (random, regular, systematic) on the estimation of mean seed production at the plot scale. One hundred quadrats were collected in 0.25 ha beech plots and considered as a reference for evaluating the different sampling schemes and sampling sizes. Measurements were performed in October (three plots), which represented the peak of seed fall, and November (two plots). Results indicate that about 25 randomly located measurements allowed to characterize plot-level mean seed production with an acceptable error below 20%, regardless of the different mean seed production observed between the studied plots and the sampling periods. If the 25 sampling points are arranged in a grid, the obtained mean estimates are within the confidence interval of the reference plot-level values. © 2021 Elsevier B.V.

• Key message: Litter trap is considered the most effective method to quantify seed production, but it is expensive and time-consuming. Counting fallen seeds using a quadrat placed on the ground yields comparable estimates to the litter traps. Ground quadrat estimates derived from either visual counting in the field or image counting from quadrat photographs are comparable, with the latter being also robust in terms of user sensitivity. • Context: Accurate estimates of forest seed production are central for a wide range of ecological studies. As reference methods such as litter traps (LT) are cost- and time-consuming, there is a need of fast, reliable, and low-cost tools to quantify this variable in the field. • Aims: To test two indirect methods, which consist of counting the seeds fallen in quadrats. • Methods: The trial was performed in three broadleaved (beech, chestnut, and Turkey oak) tree species. Seeds are either manually counted in quadrats placed at the ground (GQ) or from images acquired in the same quadrats (IQ) and then compared against LT measurements. • Results: GQ and IQ provide fast and reliable estimates of seeds in both oak and chestnut. In particular, IQ is robust in terms of user sensitivity and potentially enables automation in the process of seed monitoring. A null-mast year in beech hindered validation of quadrats in beech. • Conclusion: Quadrat counting is a powerful tool to estimate forest seed production. We recommend using quadrats and LT to cross-calibrate the two methods in case of estimating seed biomass. Quadrats could then be used more routinely on account of their faster and simpler procedure to obtain measurements at more spatially extensive scales. © 2021, The Author(s).

Masting is a complex mechanism which is mainly driven by a combination of internal plant resources and climatic conditions. While the driving role of climate in masting is being intensively studied, the interplay among climate, seed production, vegetation growth and phenology still needs further investigation. The objectives of this study were to identify the climatic determinants of different levels of seed production and of NDVI-based vegetation growth and phenology in European beech, and to evaluate if exists a trade-off between these two plant processes. To answer these questions, we used a 25-year-long dataset of beech seed production. We exploited the concept of ecological niche assuming that a mast year can be modeled like a species with variable preferences for different resources, which are the underlying annual climatic conditions; we performed an Ecological Niche Factor Analysis (ENFA), a presence-only modeling tool conventionally used in zoology and botany, and used seasonal (spring, summer, autumn) Standardized Precipitation-Evaporation Index (SPEI) observations, considering the current year (y−0), and up to one (y−1) and two (y−2) years before the masting event. For analyzing the role of vegetation growth and phenology, we used seasonal Normalized Difference Vegetation Index (NDVI) values and associated NDVI-based phenological metrics derived from Landsat imagery. Results indicated the driving role of climate for masting, especially in VHSP years. A moist summer and dry spring at y−2 and a dry summer at y−1 represented the main driving climatic conditions for masting; while a moist spring during the observation year represented the key condition for triggering higher intensities of seed production. Summer NDVI at y−0 and y−1 represented the variables discriminating best between masting and non-masting years and resulted as driven by opposite summer climatic conditions than seed production, thus indicating a trade-off between seed production and vegetation phenology. We concluded that reproduction and vegetation growth act as two different climate-dependent plant responses in beech, in a way that certain conditions through the years promote mast seeding and the opposite conditions favor vegetation growth. The understanding of climate-growth-masting relationships represents indispensable knowledge for providing a holistic view of masting mechanisms and developing adaptive forest management strategies in this species. © 2020