Herbivores are constitutive elements of most terrestrial ecosystems. Understanding effects of herbivory on ecosystem dynamics is thus a major, albeit challenging task in community ecology. Effects of mammals on plant communities are typically explored by comparing plant densities or diversity in exclosure experiments. This might over-estimate long-term herbivore effects at community levels as early life stage mortality is driven by a multitude of factors. Addressing these challenges, we established a set of 100 pairs of ungulate exclosures and unfenced control plots (25 m2) in mixed montane forests in the Alps in 1989 covering a forest area of 90 km2. Investigations ran until 2013. Analogous to the gap-maker–gap-filler approach, dynamically recording the height of the largest trees per tree species in paired plots with and without exclosures might allow for assessing herbivore impacts on those individuals with a high probability of attaining reproductive stages. We thus tested if recording maximum heights of regenerating trees would better reflect effects of ungulate herbivory on long-term dynamics of tree regeneration than recording of stem density, and if species dominance patterns would shift over time. For quantifying the effects of ungulate herbivory simultaneously at community and species level we used principle response curves (PRC). PRCs yielded traceable results both at community and species level. Trajectories of maximum heights yielded significant results contrary to trajectories of total stem density. Response patterns of tree species were not uniform over time: e.g., both Norway spruce and European larch switched in their response to fencing. Fencing explained about 3% of the variance of maximum tree heights after nine years but increased to about 10% after 24 years thus confirming the importance of long-term surveys. Maximum height dynamics of tree species, addressed in our study, can thus reflect local dominance of tree species via asymmetric plant competition. Such effects, both within and among forest patches, can accrue over time shaping forest structure and composition.
Mast seeding, the synchronized interannual variation in seed production of trees, is a well‐known bottom‐up driver for population densities of granivorous forest rodents. Such demographic effects also affect habitat preferences of the animals: After large seed production events, reduced habitat selectivity can lead to spillover from forest patches into adjacent alpine meadows or clear‐cuts, as has been reported for human‐impacted forests. In unmanaged, primeval forests, however, gaps created by natural disturbances are typical elements, yet it is unclear whether the same spillover dynamics occur under natural conditions. To determine whether annual variation in seed production drives spillover effects in naturally formed gaps, we used 14 years of small mammal trapping data combined with seed trap data to estimate population densities of Apodemus spp. mice and bank voles (Myodes glareolus) on 5 forest sites with differing disturbance history. The study sites, located in a forest dominated by European beech (Fagus sylvatica), Norway spruce (Picea abies), and silver fir (Abies alba), consisted of two primeval forest sites with small canopy gaps, two sites with larger gaps (after an avalanche event and a windthrow event), and a managed forest stand with closed canopy as a control. Hierarchical Bayesian N‐mixture models revealed a strong influence of seed rain on small rodent abundance, which were site‐specific for M. glareolus but not for Apodemus spp. Following years of moderate or low seed crop, M. glareolus avoided open habitat patches but colonized those habitats in large numbers after full mast events, suggesting that spillover events also occur in unmanaged forests, but not in all small rodents. The species‐ and site‐specific characteristics of local density responding to food availability have potentially long‐lasting effects on forest gap regeneration dynamics and should be addressed in future studies.
Different species-specific traits of woody plant species, feeding preferences of herbivores together with resulting effects on plant competition are expected to translate into different plant community structures and expressions of biodiversity. We studied the diversity of woody plant species (trees and shrubs) and structural diversity of forest trees, using a 30-year and an 18-year dataset of ungulate exclosure-control plot pairs in a mixed alpine forest community in Austria. We surveyed the tallest individuals per tree species and plot and analyzed the collective of top-height individuals per plot pair. Incidence data for exclosure and control plots were aggregated. Comparing species diversity and diversity of height classes on the plots throughout time, we calculated diversity profiles based on Hill numbers. Diversity of top height individuals and structural diversity, expressed by height classes, were two diversity aspects that differed between exclosures and control plots. Other diversity estimates of woody plant species showed huge variation without significant differences between plots. Height growth was significantly suppressed by ungulate herbivory. Effects of ungulate herbivores in forest ecosystems are highly complex and context-dependent and thus not reducible to simple top-down forces. Long-term surveys provide data that reflect “ultimate” effects of herbivory interacting with other drivers of community dynamics.
Food-hoarding granivores act as both predators and dispersers of plant seeds, resulting in facultative species interactions along a mutualism–antagonism continuum. The position along this continuum is determined by the positive and negative interactions that vary with the ratio between seed availability and animal abundance, particularly for mast-seeding species with interannual variation and spatial synchrony of seed production. Empirical data on the entire fate of seeds up to germination and the influence of rodents on seed survival is rare, resulting in a lack of consensus on their position along the mutualism–antagonism continuum. Here, we quantified annual seed rain and rodent abundance in an old-growth European beech forest and tracked 639 beechnuts to the seedling stage with 84% of seeds successfully located. Over 4 study years that covered the range of seed-to-rodent ratios, not a single seed successfully germinated after dispersal, illustrating a predominantly antagonistic interaction between rodents and seeds of European beech. Therefore, our findings do not support the predator dispersal hypothesis and partially contradict the predator satiation hypothesis, as the highest number of germinants and intact seeds were found in situ after an intermediate seed crop, not a bumper crop. Our results underline the necessity to track seeds up to germination.
The concept of landscape connectivity involves species movement between habitat patches influenced by landscape features. It encompasses structural and functional connectivity as well as species-specific considerations. Structural connectivity analyzes spatial patterns of landscapes, while functional connectivity considers the response of organisms to the landscape. Evaluating habitats for connectivity requires accounting for their spatial and temporal variations. Temporal connectivity—often overlooked—is particularly essential for long-term population viability. Conservation planning should therefore integrate monitoring and assessment measures to achieve connectivity objectives in dynamic landscapes affected by land use and climate change. Measuring landscape connectivity considers landscape composition, structure, and heterogeneity as well as the presence of barriers, each varying among species and scales. Assessing connectivity across scales requires considering biological levels of organization from genetic flow to community processes. Modeling connectivity is complex and incorporates patch- and landscape-based approaches. Patch-based models focus on attributes of habitat patches, while landscape-based models consider movement behavior and resistance surfaces. Landscape connectivity research has expanded rapidly in recent decades, and its conceptual foundations are evolving. Recent advances integrate metapopulation dynamics with habitat configuration and movement behavior. Traditional static models are being replaced with dynamic models considering temporal variations in landscape attributes. Further technological advancements such as remote sensing and climate simulators allow more accurate representations of dynamic landscapes, promoting ecosystem understanding and supporting conservation planning.