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Accurate individual tree segmentation (ITS) is fundamental to forest management and to the studies of forest ecosystem. Unmanned Aerial Vehicle Light Detection and Ranging (UAV-LiDAR) shows advantages for ITS and tree height estimation at stand and landscape scale. However, dense deciduous forests with tightly interlocked tree crowns challenge the performance for ITS. Available LiDAR points through tree crown and appropriate algorithm are expected to attack the problem. In this study, a new UAV-LiDAR dataset that fused leaf-off and leaf-on point cloud (FULD) was introduced to assess the synergetic benefits for ITS and tree height estimation by comparing different types of segmentation algorithms (i.e., watershed segmentation, point cloud segmentation and layer stacking segmentation) in the dense deciduous forests of Northeast China. Field validation was conducted in the four typical stands, including mixed broadleaved forest (MBF), Mongolian oak forest (MOF), mixed broadleaf-conifer forest (MBCF) and larch plantation forest (LPF). The results showed that the combination of FULD and the layer stacking segmentation (LSS) algorithm produced the highest accuracies across all forest types (F-score: 0.70 to 0.85). The FULD also showed a better performance on tree height estimation, with a root mean square error (RMSE) of 1.54 m at individual level. Compared with using the leaf-on dataset solely, the RMSE of tree height estimation was reduced by 0.22 to 0.27 m, and 12.3% more trees were correctly segmented by the FULD, which are mainly contributed by improved detection rate at nearly all DBH levels and by improved detection accuracy at low DBH levels. The improvements are attributed to abundant points from the bole to the treetop of FULD, as well as each layer point being included for segmentation by LSS algorithm. These findings provide useful insights to guide the application of FULD when more multi-temporal LiDAR data are available in future.

Summary Within the same forest stand, temperate deciduous trees generally exhibit a distinct pattern in leaf‐out timing, with some species flushing earlier than other species. This study aimed to explain the timing of leaf‐out of various temperate tree species in relation to the risk of freezing damage to leaves. We combined long‐term series of leaf‐out date (14–32 years) of five temperate tree species located in both low and high elevations in Switzerland, daily minimum temperatures recorded at the same sites and species‐specific freezing resistance (LT50) of emerging leaves. We calculated temperature safety margins (the temperature difference between absolute minimum temperature during leaf‐out and species‐specific LT50 values), and date safety margins (time lag between the last day when temperature falls below species‐specific LT50 values and the date of leaf‐out). Leaf‐out occurred when the probability to encounter freezing damage approaches zero, irrespective of climatic conditions (low vs. high elevation) and species (early‐ and late‐flushing species). In other words, trees leaf out precisely at the beginning of the probabilistically safe period. Interestingly, the temperature safety margins did not differ significantly between low and high elevation. Yet, the date safety margin was smaller at high elevation, presumably due to a faster increase in temperature during the leaf‐out period at high elevation. When species‐specific freezing resistance is taken into account, the time of leaf‐out converges among species towards a marginal risk of freezing damage. Thus, leaf‐out time has likely evolved in a way that the risk of freezing damage is minimized over a large spectrum of climatic conditions. Species with a small safety margin against freezing temperature, like Fagus sylvatica, appear to employ photoperiod co‐control of spring phenology, whereas species with a large safety margin depend largely on temperature for the right timing of leaf‐out. Our results offer a new avenue to explain the differences in leaf‐out timing among co‐occurring tree species. They further suggest that in a warming climate, tree species can expand their distribution range to the extent their phenology matches the stochasticity of freezing temperatures in spring. Lay Summary

Aims To develop forest vegetation classification at the level of alliances and associations across the Hyrcanian ecoregion, Northern Iran, and to explore the effects of main environmental and geographic gradients on their distribution. Location Hyrcanian ecoregion, Northern Iran. Methods A database of 1,597 vegetation plots of mostly 400 m2 in size with a total of 802 vascular plant taxa was established, covering the whole geographic range of the Hyrcanian forests at altitudes ranging from −22 to 2,850 m a. s. l. An expert system was developed for automatic classification of vegetation plots into alliances and associations. Detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) were used to determine the most important environmental and geographic gradients affecting species composition. Results Twenty‐one associations and seven alliances of these forests, belonging to five orders and four classes, were defined. Among them, eleven associations and five alliances were described as new syntaxa. Alnion glutinosae and Smilaco excelsae‐Alnion barbatae, both distributed in the lowland belt, include swamp and wet forests, respectively. Parrotio persicae‐Carpinion betuli and Alnion subcordatae are respectively mesic and wet forests of the submontane belt. Solano kieseritzkii‐Fagion orientalis is a mesic beech forest in the montane belt, and finally, Quercion macrantherae, an open oak forest, and Centaureo hyrcanicae‐Carpinion orientalis, a dry hornbeam forest, occur in the upper‐montane belt. DCA and CCA analyses showed that the distribution of these alliances and associations is mainly related to altitude and mean annual temperature. Conclusions Based on our results and comparison between the Hyrcanian and European forests, we propose a new, comprehensive syntaxonomic scheme for the Hyrcanian forests, supported by a classification expert system. Unlike previous studies, we linked the classification system to that of EuroVegChecklist because, though this area is outside of Europe, its vegetation is very similar to that of the European temperate forest vegetation. The Hyrcanian ecoregion in Northern Iran is an isolated area with a well‐developed biome of temperate broad‐leaved deciduous forest, containing many relict species. We provide the first comprehensive classification of these forests into alliances and associations across the whole ecoregion, based on a new database of 1,597 vegetation plots, and relate individual vegetation types to altitude, climate and soil conditions.

The climate sensitivity of forest ecosystem woody productivity (ANPPstem ) influences carbon cycle responses to climate change. For the first time, we combined long-term annual growth and forest census data of a diverse temperate broadleaf deciduous forest, seeking to resolve whether ANPPstem is primarily moisture- or energy-limited and whether climate sensitivity has changed in recent decades characterised by more mesic conditions and elevated CO₂. We analysed tree-ring chronologies across 109 yr of monthly climatic variation (1901–2009) for 14 species representing 97% of ANPPstem in a 25.6 ha plot in northern Virginia, USA. Radial growth of most species and ecosystem-level ANPPstem responded positively to cool, moist growing season conditions, but the same conditions in the previous May–July were associated with reduced growth. In recent decades (1980–2009), responses were more variable and, on average, weaker. Our results indicated that woody productivity is primarily limited by current growing season moisture, as opposed to temperature or sunlight, but additional complexity in climate sensitivity may reflect the use of stored carbohydrate reserves. Overall, while such forests currently display limited moisture sensitivity, their woody productivity is likely to decline under projected hotter and potentially drier growing season conditions.

A stable below-canopy microclimate of forests is essential for their biodiversity and ecosystem functionality. Forest management necessarily modifies the buffering capacity of woodlands. However, the specific effects of different forestry treatments on site conditions, the temporal recovery after the harvests, and the reason for the contrasts between treatments are still poorly understood. The effects of four different forestry treatments (clear-cutting, retention tree group, preparation cutting, and gap-cutting) on microclimatic variables were studied within a field experiment in a managed oak-dominated stand in Hungary, before (2014) and after (2015–2017) the interventions by complete block design with six replicates. From the first post-treatment year, clear-cuts differed the most from the uncut control due to the increased irradiance and heat load. Means and variability of air and soil temperature increased, air became dryer along with higher soil moisture levels. Retention tree groups could effectively ameliorate the extreme temperatures but not the mean values. Preparation cutting induced slight changes from the original buffered and humid forest microclimate. Despite the substantially more incoming light, gap-cutting could retain the cool and humid air conditions and showed the highest increase in soil moisture after the interventions. For most microclimate variables, we could not observe any obvious trend within 3 yr. However, soil temperature variability decreased with time in clear-cuts, while soil moisture difference continuously increased in gap- and clear-cuts. Based on multivariate analyses, the treatments separated significantly based mainly on the temperature maxima and variability. We found that (1) the effect sizes among treatment levels were consistent throughout the years, (2) the climatic recovery time for variables appears to be far more than 3 yr, and (3) the applied silvicultural methods diverged mainly among the temperature maxima. Based on our study, the spatially heterogeneous and fine-scaled treatments of continuous cover forestry (gap-cutting, selection systems) are recommended. By applying these practices, the essential structural elements creating buffered microclimate could be more successfully maintained. Thus, forestry interventions could induce less pronounced alterations in environmental conditions for forest-dwelling organism groups.

As climate change drives increased drought in many forested regions, mechanistic understanding of the factors conferring drought tolerance in trees is increasingly important. The dendrochronological record provides a window through which we can understand how tree size and traits shape growth responses to droughts. We analyzed tree-ring records for 12 species in a broadleaf deciduous forest in Virginia (USA) to test hypotheses for how tree height, microenvironment characteristics, and species’ traits shaped drought responses across the three strongest regional droughts over a 60-yr period. Drought tolerance (resistance, recovery, and resilience) decreased with tree height, which was strongly correlated with exposure to higher solar radiation and evaporative demand. The potentially greater rooting volume of larger trees did not confer a resistance advantage, but marginally increased recovery and resilience, in sites with low topographic wetness index. Drought tolerance was greater among species whose leaves lost turgor (wilted) at more negative water potentials and experienced less shrinkage upon desiccation. The tree-ring record reveals that tree height and leaf drought tolerance traits influenced growth responses during and after significant droughts in the meteorological record. As climate change-induced droughts intensify, tall trees with drought-sensitive leaves will be most vulnerable to immediate and longer-term growth reductions.

Invasive fungal pathogens are an increasing problem globally and can cause strong effects on forest ecosystems. In this study, we contrast vegetation surveys in eutrophic elm (Ulmus glabra) and ash (Fraxinus excelsior) forests in southern Sweden, conducted just prior to the arrival of Dutch elm disease (DED) in 1989, and then again in 2021, several years after ash dieback (ADB) began. At the sample plot scale, species richness (α–diversity) of the upper tree layer strongly decreased from 1989 to 2021, and the mean cover of elm decreased from 27 to 1% and of ash from 29 to 13%. In the lower tree and shrub layers, elm and ash were replaced by other, mainly shade-tolerant, tree species. The cover and richness of the shrub layer increased in previously elm-dominated stands but not in ash-dominated stands. The extensive loss of canopy cover in elm stands caused a larger change in upper tree layer species composition and increased compositional variability (β-diversity) between plots when compared to the ash stands. The direction of the changes in tree layer composition between the surveys varied with soil moisture and nutrient availability. While beech increased in less eutrophic plots, more nutrient-rich plots changed toward hornbeam or small-leaved lime, and wetter plots turned toward alder and bird cherry. Hence, our results indicate increased compositional diversity and alternative successional pathways for community reorganization following DED and ADB. Future research will reveal if these pathways will later merge or further split.

A major goal in ecology is to understand mechanisms that increase invasion success of exotic species. A recent hypothesis implicates altered species interactions resulting from ungulate herbivore overabundance as a key cause of exotic plant domination. To test this hypothesis, we maintained an experimental demography deer exclusion study for 6 y in a forest where the native ungulate Odocoileus virginianus (white-tailed deer) is overabundant and Alliaria petiolata (garlic mustard) is aggressively invading. Because population growth is multiplicative across time, we introduce metrics that correctly integrate experimental effects across treatment years, the cumulative population growth rate, λc, and its geometric mean, λper-year, the time-averaged annual population growth rate. We determined λc and λper-year of the invader and of a common native, Trillium erectum. Our results conclusively demonstrate that deer are required for the success of Alliaria; its projected population trajectory shifted from explosive growth in the presence of deer (λper-year = 1.33) to decline toward extinction where deer are excluded (λper-year = 0.88). In contrast, Trillium's λper-year was suppressed in the presence of deer relative to deer exclusion (λper-year = 1.04 vs. 1.20, respectively). Retrospective sensitivity analyses revealed that the largest negative effect of deer exclusion on Alliaria came from rosette transitions, whereas the largest positive effect on Trillium came from reproductive transitions. Deer exclusion lowered Alliaria density while increasing Trillium density. Our results provide definitive experimental support that interactions with overabundant ungulates enhance demographic success of invaders and depress natives' success, with broad implications for biodiversity and ecosystem function worldwide.

Recent studies have shown that complex species interactions can regulate above- and belowground processes in terrestrial systems. Ungulate herbivory and invasive species are known to have strong effects on plant communities in some systems, but their impacts on soil biota and belowground processes are lesser known. Growing evidence suggests white-tailed deer (Odocoileus virginianus) and invasive plants facilitate increased abundance of exotic earthworms in temperate forests of the eastern United States. We conducted an experimental study that manipulated deer access and the presence of an invasive understory shrub in an eastern deciduous forest of southwestern Ohio, USA, from 2013 to 2017. Earthworm density and biomass, and standing litter biomass were measured in five paired deer access and exclosure plots, each with a split-plot removal of Amur honeysuckle (Lonicera maackii). Earthworm density declined in response to the experimental exclusion of deer, with earthworm density decreasing over time in the deer exclosure plots relative to deer access plots. Deer exclusion produced greater variation in earthworm species composition relative to access plots. Multivariate analyses indicated that larger earthworms in the genus Lumbricus were associated with deer exclosure plots, while smaller endogeic species were ubiquitous in both treatments. Standing litter biomass decreased over time in the deer-access plots. In contrast, honeysuckle removal had little effect on earthworm density and standing litter biomass. There was an interaction between deer and honeysuckle treatments on earthworm biomass, with honeysuckle removal reducing earthworm biomass when deer were excluded. Our results demonstrate strong effects of herbivores on invasive earthworms and ecosystem processes, but indicate a weaker influence of invasive shrubs. Further, our findings suggest that the effects of deer overabundance in forest ecosystems are potentially reversible with long-term intervention.

Forests play an important role in climate regulation due to carbon sequestration. However, a deeper understanding of forest carbon flux dynamics is often missing due to a lack of information about forest structure and species composition, especially for non-even-aged and species-mixed forests. In this study, we integrated field inventory data of a species-mixed deciduous forest in Germany into an individual-based forest model to investigate daily carbon fluxes and to examine the role of tree size and species composition for stand productivity. This approach enables to reproduce daily carbon fluxes derived from eddy covariance measurements (R2 of 0.82 for gross primary productivity and 0.77 for ecosystem respiration). While medium-sized trees (stem diameter 30–60 cm) account for the largest share (66%) of total productivity at the study site, small (0–30 cm) and large trees (>60 cm) contribute less with 8.3% and 25.5% respectively. Simulation experiments indicate that vertical stand structure and shading influence forest productivity more than species composition. Hence, it is important to incorporate small-scale information about forest stand structure into modelling studies to decrease uncertainties of carbon dynamic predictions.