Browsing Pressure Research Articles

Tree growth‐forms reveal dominant browsers shaping the vegetation

Abstract Plants adopt particular growth‐forms when they are exposed to extreme environmental conditions. In this study, we describe a unique woody plant growth‐form induced by large mammalian herbivores and discuss that this growth‐form could have evolved as a strategy for escaping the browser zone in herbivore driven ecosystems. We analysed responses of key architectural and morphological attributes (branching and thorn density, tree dimensions, presence of flowers and fruits) of three Eurasian spiny tree species (Malus sylvestris, Prunus cerasifera and Pyrus communis) to different levels of browsing by large herbivores in the temperate Białowieża Forest, Poland. Under high browsing pressure, studied trees displayed two distinct forms of the crown: a bottom sterile part developing into a densely branched structure with high density of thorns (‘cage‐form’), and an upper reproductive part that escaped from herbivore control (‘escaped‐form’). The size of cage‐form influenced the feeding behaviour of red deer (Cervus elaphus) by increasing the time deer spend foraging and increasing the bite rate. The height at which cages started to escape and their diameter matched with foraging reach of red deer. Synthesis. We argue that the frequency and cage dimensions of this woody growth‐form in the landscape could inform on the type and intensity of recent herbivory. Moreover, its distinctive inducibility suggests that this growth‐form did not emerge recently under anthropogenic pressure but could be the legacy of ancient herbivory effects. Observational evidence suggests that this growth‐form emerged in several herbivore‐driven systems around the globe and may be used to identify the dominant herbivores that control vegetation structure in these ecosystems. Read the free Plain Language Summary for this article on the Journal blog.

Deer in the agriculture-forest matrix: Interacting effects of land uses on browsing pressure on trees

Cervids inhabit mosaic landscapes that are strongly influenced by human land use and actions. Forestry and agriculture influence both the foodscape and the risk landscape, thus influencing how cervids respond to resource-risk trade-offs. Furthermore, these land use types interact to shape cervid use of the landscape. For example, cervids can forage in open agricultural fields providing high quality forage during the night, while using nearby forest patches to reduce risk during the day. Hence, actions conducted in one of these land use types will likely influence the use of the other by cervids. In this study, we investigate how agricultural and silvicultural practices interact to shape cervid use of the forest and their impact on commercially valuable trees. More specifically, we investigate the effect of growing nutrient dense crops in mixed forest-agriculture landscapes. We hypothesized that the sowing of such crops may not only attract cervids to the agricultural parts of the landscape, but also lead to increased browsing levels in adjacent forest areas where cervids seek cover. Moreover, such high quality crops may not only influence the number of animals, but also increase the intake rate of fibrous tree saplings of individual cervids to nutritionally balance the high input of easily-digestible macronutrients from the crops. We tested these ideas by conducting a large landscape-scale experiment where we manipulated the foodscape by growing two crop types on 28 fields (14 fields with high-quality oat versus 14 fields with lower-quality grass) and measuring cervid use of tree saplings in the forest patches adjacent these fields. We also investigated how forage availability in the forest patches and the distance from field edge affected browsing on tree saplings. Finally, we compared the effects of crop type and forest forage availability on the browsing pressure on coniferous versus deciduous tree species. We studied this in a multispecies ungulate system, where four cervid species occur sympatrically. We show that the type of crop sown influenced browsing pressure on tree saplings in nearby forests, with higher browsing levels in forests surrounding oat compared to grass fields. Moreover, browsing pressure in these forests was influenced by the availability of young trees in the forest stands, where more young trees led to reduced browsing pressure on individual saplings. Our study emphasizes that human practices that shape features of the foodscape in one land use type will influence the impact cervids have in another land use type. Hence, our study highlights the importance of considering the actions and impacts of different land uses simultaneously for predicting cervid foraging patterns in mixed forest-agriculture landscapes.

Revegetated riparian areas are dominated by weeds, and lack structural diversity and natural recruitment: lessons for restoration practice

Riparian areas can be highly biodiverse and provide critical ecosystem services. However, they are frequently subject to anthropogenic impacts such as land clearing, agricultural use, and urban development. Restoration of riparian areas via revegetation commonly improves the health of waterways and surrounding areas, but vegetation outcomes are rarely assessed. Our study compared 10 to 14‐year‐old revegetation to remnant vegetation in riparian areas of south‐eastern Australia to determine if species composition, vegetation structure, and ecosystem function (plant recruitment) differed. We also assessed if the amount of surrounding native vegetation, browsers (including deer, rabbits, and macropods), or soil characteristics influenced native woody plant recruitment at revegetated sites. While native and exotic woody species richness did not differ, native woody plants were less abundant and exotic woody plants twice as abundant at revegetated sites. The ground layer of revegetated sites was dominated by weeds, whereas remnant sites largely comprised native herbaceous plants and leaf litter. Tree heights and tree canopy cover were similar in revegetated and remnant areas, although shrubs and ferns were lacking in revegetation. Native woody plant recruitment was lower at revegetated sites and was negatively associated with browser presence. Our results suggest that while revegetation may have similar species richness and tree cover as remnant areas, weeds often dominate and important structural components such as shrubs and ferns, and ecological processes such as plant recruitment, are lacking. Better consideration of all vegetation strata, reduced browsing pressure and weed control are likely to achieve better revegetation outcomes.

Oak regeneration facing deer browsing: Can competition between saplings offset the diversion effect? A simulation experiment

The constraint caused by wild ungulates on forest regeneration is increasing worldwide. Hypotheses for plant association effects predict that species susceptible to herbivory can gain protection from other neighbouring plant species. In theory, such interactions could help limit the impact of browsing on the regeneration of specific tree species. However, the presence of neighbouring species can also result in increasing competition for resources between species. The resultant effects on forest regeneration of these interactions, both positive (protection against herbivores) and negative (inter-specific competition) are still unclear.To gain insight, we coupled models of browsing by roe deer (Capreolus capreolus) and of forest dynamics to simulate trajectories of oak (Quercus petraea (Matt.) Liebl.) regeneration admixed with species of contrasted palatability and growth rate under different scenarios of browsing pressure and initial sapling density. We also investigated how releasing oak saplings from all or specific neighbours during the simulation affect regeneration.We found that admixed species composition had a relatively weak effect on the density of oak recruits, but a strong effect on the duration of the regeneration phase. Oak regenerated faster when admixed with species of intermediate growth and low palatability (Fagus sylvatica) than with species of fast growth and high palatability (Carpinus betulus L.), except at intermediate sapling density and high browsing pressure where we found the opposite. Releasing oak from all competitors was most effective in promoting oak regeneration when admixed with both species together, although the benefit of competition release was much weaker at high browsing pressure. Lastly, we found that at low initial sapling density (i.e., 10 saplings/m2), oak regeneration was driven only by browsing and the effect of admixing species became negligible.Our study showed that admixing oak with palatable neighbours impedes rather than improves oak regeneration due to increased competition for resources. As such, we suggest that the benefits of herbivore diversion can be off-set by increased inter-specific competition.

Drivers of tree regeneration in coniferous monocultures during conversion to mixed forests in Central Europe – Implications for forest restoration management

In Central Europe, anthropogenic coniferous monocultures have been subject to conversion to more diverse mixed forests since the 1990s, however, they are still abundant across many forest landscapes. Artificial and natural tree regeneration both play a key role during conversion by determining the species composition and structure of the future forests. Many abiotic and biotic factors can potentially influence the regeneration process and its specific combinations or interactions may be different among tree species and its developmental stages. Here, we aimed to identify and quantify the effect of the most important drivers on the density of the most abundant regenerating tree species (i.e., Norway spruce and European beech), as well as on species and structural diversity of the tree regeneration. We studied tree regeneration in four former monospecific coniferous stand types (i.e., Norway spruce, Scots pine, European larch, and Douglas fir) in Southwest Germany that have been under conversion to mixed forests since the 1990s. We sampled tree regeneration in four growth height classes together with a variety of potentially influencing factors on 108 sampling plots and applied multivariate analyses. We identified light availability in the understorey, stand structural attributes, browsing pressure, and diaspore source abundance as the most important factors for the density and diversity of tree regeneration. Particularly, we revealed species-specific differences in drivers of regeneration density. While spruce profited from increasing light availability and decreasing stand basal area, beech benefited either from a minor reduction or more strikingly from an increase in overstorey density. Increasing diaspore source abundance positively and a high browsing pressure negatively affected both species equally. Our results suggest that humus and topsoil properties were modified during conversion, probably due to changes in tree species composition and silvicultural activities. The species and structural diversity of the tree regeneration benefitted from increasing light availability, decreasing stand basal area, and a low to moderate browsing pressure. We conclude that forest managers may carefully equilibrate among the regulation of overstorey cover, stand basal area, and browsing pressure to fulfil the objectives of forest conversion, i.e., establishing and safeguarding a diverse tree regeneration to promote the development of mature mixed forests in the future.

Harvest block aggregation as a driver of intensive moose browsing pressure on hardwood regeneration in a temperate forest

In many regions of the world, browsing pressure by ungulates has increased over the last century, often reaching unsustainable levels. Overbrowsing by ungulates is threatening biodiversity and causing delays in the establishment of forest regeneration. Moose (Alces alces) is considered as an ecological engineer, as it can cause severe impacts on stand structure, composition, quality, and succession through overbrowsing. In conifer-dominated stands, moose browsing pressure has been shown to vary with the distribution of forage and cover patches across the landscape, but such spatial relationships have not been explored in hardwood-dominated landscapes, where intensive forest management can create clustered patches of good-quality forage and cover, ideal conditions for wintering moose. We investigated the influence of local and landscape variables on browsing by North American moose (Alces alces americana) on commercial and non-commercial woody regeneration in an intensively managed forest landscape of northwestern New Brunswick, Canada. We predicted that browsing pressure at the local scale (vegetation sampling plot) would increase with moose winter counts in the surrounding landscape (1 to 3 km radius). We modeled moose counts as a function of environmental features and road density, using a quasi-Poisson regression. According to an aerial survey (25 km2 sampling plots), moose counts increased with the proportion of forage patches (disturbed stands < 20 years old), whereas they decreased with the proportion of mixed mature stands > 80 years old, which offer both forage and cover. We extrapolated this model to a 2770 km2 study area and used predicted counts as an independent variable to assess its influence on browsing pressure. In 720 plots of 4.15 m2, browsing pressure was quantified using occurrence (i.e. proportion of seedlings/saplings with at least one twig browsed during the previous winter) and intensity (i.e. mean proportion of twigs browsed per stem) on commercial and non-commercial tree seedlings and saplings. We explained variation in occurrence and intensity using zero-inflated models with snow precipitation and local vegetation variables as covariates. Browsing pressure increased with predicted moose counts within a 3 km radius, with snow precipitation and with the proportion of coniferous trees in the canopy, whereas it decreased with sapling density. These results highlight the importance of seedlings and saplings in moose winter diet and confirm the attraction of moose toward recent (<20-year-old) harvest blocks during winter. Therefore, we should avoid aggregating harvest blocks to decrease their attractiveness to moose and browsing pressure on natural hardwood regeneration.

The presence of wolves leads to spatial differentiation in deer browsing pressure on forest regeneration

With the recent return of large carnivores to forest ecosystems, an important issue for forest owners and managers is how large predators influence the behaviour of their natural prey and, consequently, cervid browsing pressure on forest regeneration. To investigate this issue, we analysed deer pressure on Scots pine and European beech plantations in northern Poland's ecosystems with and without permanent wolf populations. Two characteristics were used to describe deer browsing patterns in plantations: distance from the forest edge (spatial pattern of browsing) and number of saplings browsed (browsing intensity). Beech saplings were more intensively browsed by deer compared to pine saplings. In a forest ecosystem not inhabited by wolves, spatial variation in browsing patterns on small-sized beech plantations was the same between the edge and the center. In contrast, browsing pressure by deer was greater at the edges on large-sized pine plantations. The presence of wolves reduced deer browsing on beech and increased browsing on pine saplings. In addition, deer foraging behaviour changed in large-sized pine plantations, and browsing pressure increased only in the central areas of the plantations. We assume that the presence of wolves in a forest landscape is an important factor that alters browsing pressure on the youngest stands and their spatial pattern, and that this may be a major factor in stand regeneration, especially in small forest patches.

Growth response and survival of American beech, yellow birch, and sugar maple regeneration to partial harvest

In northern hardwood stands, silvicultural treatments aimed at regenerating sugar maple and yellow birch, such as site preparation and mechanical control of American beech saplings, often results in the formation of dense thickets of beech regeneration. The objective of this study was to compare the growth and survival of beech regeneration originating from stump sprouts, root suckers and seedlings, with yellow birch and sugar maple regeneration in partially harvested stands under cervid browsing pressure. Over an eight-year period, 997 individuals of the three species were monitored in three northern hardwood stands dominated by sugar maple, yellow birch, and American beech in Quebec, Canada. Each stand included replicates of four partial harvesting treatments and a control, providing a gradient of light and competition conditions. Beech stump sprouts and root suckers had higher growth and survival probabilities than seedlings of all three species under all light and competition conditions. Results suggest that treatments triggering beech sprouts should be applied with caution and under conditions that ensure sufficient survival and development of desired species. This study also provides evidence that selective browsing by cervids could be an important overriding factor exacerbating the effect of beech competition on yellow birch regeneration.

Heavy browsing pressure by moose (Alces alces) can interfere with the objectives of ecosystem-based forest management

Forest management can enhance the ecosystem services forests provide, but it can also alter the capacity of forests to provide those services in the future. Ecosystem-based management has been widely implemented to mitigate the deterioration of forest ecosystems by reducing dissimilarities between managed and natural forests. Ecosystem-based management objectives generally aim to maintain community composition, stand structure, ecosystem functions and services. Heavy browsing pressure by ungulates at densities unrepresentative of past abundance could interfere with these objectives by chronically and selectively browsing tree regeneration following logging.Using a browsing exclusion experiment, we investigated how the effects of different moose browsing pressures on tree regeneration could interfere with ecosystem management objectives in boreal forests of Quebec, Canada. Management objectives in this region include, among others, reaching targeted coniferous trees densities for commercial harvest, favouring the conservation of birches (Betula spp.) to maintain soil quality and conserving fruit-bearing trees due to their ecological significance for biodiversity. We used path analyses to differentiate direct and indirect effects of moose on seedling density and mixed models to investigate how browsing affected sapling recruitment and whether resulting densities are compatible with ecosystem management objectives.By directly reducing the overall sapling density, moose browsing indirectly increased seedlings densities of all deciduous species. This indirect effect increased with browsing pressure and favoured raspberry cover, which was positively or negatively linked to seedling density, depending on tree species. Moderate and heavier browsing pressures reduced coniferous sapling density below the ecosystem management target. Moose suppressed birch saplings so that the minimal target of 50 birches ha−1 was not met under heavier browsing pressure. Sapling density of fruit-bearing trees was low under all browsing pressures when compared to exclosures.Policy implicationsHeavy moose browsing is preventing the achievement of ecosystem management objectives in our study area. Positive indirect effects on seedling densities suggest that reducing moose density could allow released seedlings to replenish the sapling layer, but we discuss why this would not guarantee reaching ecosystem management objectives. To help forest managers evaluate and respond to challenges posed by high ungulate densities, we suggest incorporating browsing inventories in national forest surveys to monitor this chronic disturbance and modulate decisions for forest ecosystem management.

Successful forest restoration using plantation at high deer density: How neighboring vegetation drives browsing pressure and tree growth

Browsing can be an environmental stress to forest ecosystems, where the composition and structure of plant communities depend on the balance between ungulate browsing and the ability of plants to tolerate and acclimate to this stressor. Tree planting is a management tool for restoring forest whose integrity has been compromised by the intensity and repetition of browsing. This tool may be insufficient when the stress leading to forest degradation is ongoing. Balsam fir (Abies balsamea (L.) Mill.) plantations are used on Anticosti Island (Quebec, Canada) to restore winter habitat for white-tailed deer (Odocoileus virginianus Zimmermann, 1780). To promote natural regeneration and growth of planted firs that are the staple winter forage of deer, enclosures were temporarily erected and deer densities reduced to decrease browsing pressure. The stress induced by heavy browsing at the time of fence removal and the density of competitors in regenerating cutblocks, however, could compromise fir growth and forest recruitment. Our objective was to identify factors influencing browsing intensity and balsam fir’s growth following fence removal. We aimed to identify factors influencing lateral browsing intensity and number of trunk reiterations on fir at the landscape scale, including distance and size of the surrounding forests, and at the local scale through intraspecific density-dependence and interspecific associative effects. We then sought to identify the stressors most influencing balsam fir growth among browsing variables and a neighborhood crowding index (NCI). We measured browsing intensity and growth of 114 balsam firs 8 years following planting and again 2 and 4 years after dismantling two management enclosures. At the landscape scale, our results showed that when balsam fir was more than 150 m from a forest edge, the probability of browsing was close to zero and consistently a low percentage of shelter area surrounding a focal fir led to a decrease in browsing pressure. At the local scale, intensity of lateral browsing increased with the NCI of white spruce (Picea glauca (Moench) Voss). Fir growth was negatively influenced by the additive effects of browsing and NCI. Finally, our results showed that 19.7% (95 %CI: 4.5%, 42.6%) of the effect of the surrounding vegetation on growth was mediated by browsing. The success of forest ecosystem restoration through planting depends on density of neighboring tree species that can generate environmental stress by influencing growth indirectly through browsing as well as directly through competition for resources.

Large wood supports Elwha revegetation by reducing ungulate browsing

IntroductionThe increasing number of dams approaching obsolescence drives a need for knowledge about riparian restoration associated with dam removal. Restoring woody vegetation on exposed reservoir beds following dam removal is essential to stabilizing sediment, reconnecting riverine and terrestrial systems, and providing future sources of shade, nutrients, and wood. Revegetation after dam removal on many rivers can be challenging due to rapidly drying sediment, low sediment nutrient content, and heavy ungulate browse pressure. Revegetation in Elwha River restoration, the largest dam removal to date, used large woody debris (LWD) to mitigate moisture and nutrient limitation but ungulate browsing has constrained woody plant growth in many coarse sediment deposits. We evaluated potential for LWD to reduce ungulate browsing following Elwha dam removal.MethodsWe studied LWD mitigation of browsing in the largest former reservoir and a comparable valley upriver with a natural floodplain. We measured browse intensity in randomly located plots stratified by four levels of LWD extent, from no LWD to complete LWD enclosure.ResultsLWD reduced browse intensity four-fold in the former reservoir, but only in plots fully surrounded by LWD. Partial LWD enclosure provided little browse reduction. We obtained similar results in the upriver valley, where browse intensity was somewhat lower except within wood clusters. Wood-mediated browse reduction was slightly greater in the former reservoir than in the upriver valley. Protection from browse was greatest for plant species preferred by ungulates.DiscussionThese results suggest forest restoration after dam removal can be expedited by surrounding young trees with large logs. Planting within LWD clusters or placing LWD clusters in restoration sites can facilitate establishment of forest islands in strategic locations. These forest islands can support dispersal of seeds and marine derived nutrients, reconnect established forest to the river, and potentially advance restoration by decades.

Testing the response of northern white-cedar to simulated browsing: Evidence of apparent compensatory growth

The impacts of ungulate browsing on forested ecosystems largely stem from events happening at the plant scale, and plant-level responses to browsing vary according to factors including light environment and browsing intensity, frequency, and timing. Northern white-cedar (Thuja occidentalis), a slow-growing, shade tolerant, and highly browse-selected species, is experiencing regeneration problems in large part due to heavy browsing pressure by white-tailed deer (Odocoileus virginianus). Although survival and recovery following browsing are essential for cedar recruitment, we know little about the reactions of individual cedar plants to browsing. Our objective was to evaluate the impact of browsing, as emulated by clipping, on cedar height, diameter, and branch growth. We applied combinations of shading (full sun or 50% sun) and clipping timing (early growth or dormancy), intensity (apex or 50% previous year’s growth), and frequency (one or two years) to planted cedar seedlings. Partially shaded trees had systematically faster height, diameter and branch growth than trees in full sun. Clipping timing had no effect on height or branch growth but growing season clipping negatively affected diameter growth. Clipping frequency had no detectable effect on cedar growth. We observed aboveground growth compensation except under high clipping intensity. We confirm the expected browse tolerance of cedar and its high recovery potential following low to moderate browsing pressure. We underline that higher clipping intensities or longer-term browsing pressure than those we tested, which can be expected in areas with heavy, sustained browsing pressure, could negatively impact cedar’s recovery potential.

Allometric scaling models for tree saplings: Assessing potential fodder for ungulate herbivores

Knowledge of the quantity of twig dry mass (available fodder) of saplings is crucial for understanding tree recruitment under wild ungulate browsing pressure in forest ecosystems. Therefore, this study aimed to develop allometric equations for estimating twig dry mass using sapling structural data and to model the twig dry mass of saplings of various species in various size classes while quantitatively describing the variations among tree species. Sapling biomass samples were taken for six widely spread tree species in two national parks in Poland. After drying the biomass samples in the oven for 72 h at 65 °C, the data were analyzed, and models were generated using the ordinary least squares and generalized additive methods. The results confirmed our hypothesis that different tree species differ in their allometric properties. The twig dry masses of Acer pseudoplatanus and Carpinus betulus were best estimated by basal diameter, whereas those of Picea abies, Abies alba, and Fagus sylvatica were best estimated by sapling height. In contrast, the model quality for Fagus sylvatica and Sorbus aucuparia was enhanced by combining the sapling height with the mean largest crown length (MLCL). For all species, the twig dry mass distribution was linear along sapling height classes and exponentially increased with shoot diameter classes, with coniferous species exhibiting the highest correlation. On the other hand, the proportion of twig dry mass to total sapling dry mass followed the order S. aucuparia > A. alba > P. abies > F. sylvatica > C. betulus > A. pseudoplatanus. The developed allometric equations contribute significantly to the already available collection of published biomass equations. Moreover, they extremely useful in estimating the browsing potential of tree saplings falling within size ranges conducive to herbivore browsing.

Seedling growth and spinescence of three Vachellia species' populations in valleys formed by tectonic uplift

AbstractThe Southeastern Coastal Hinterland geomorphic province of South Africa is dissected by deep valleys created by tectonic uplift in the early Miocene and late Miocene/early Pliocene. We hypothesised that seedlings of Vachellia nilotica, V. tortilis and V. karroo grown from seeds from different valleys would differ in growth and spinescence because of geographical separation. Ninety seedlings (30 per species) were grown for 3 months in uniform conditions in a greenhouse. Mean height of V. nilotica and V. tortilis seedlings was greatest for the Phongolo populations and lowest for the Thukela populations. Maximum stem diameter of both species was attained by individuals from the Phongolo population. Mean spine dry mass of V. nilotica seedlings was greatest for the Phongolo population, suggesting a growth‐defence trade‐off, but did not differ for V. tortilis. Results suggested that Phongolo populations of V. nilotica and V. tortilis may be characterised by morphotypes clearly distinct from those in the other valleys. Unclear and inconsistent results for V. karroo in this study suggested subtly different morphotypes occur in each valley. We propose that distinct Phongolo morphotypes of V. nilotica and V. tortilis are explained as evolutionary adaptations to moderate fire and high browsing pressure compared to populations South of Phongolo.

Effects of understory characteristics on browsing patterns of roe deer in central European mountain forests.

Selective browsing by deer on young trees may impede the management goal of increasing forest resilience against climate change and other disturbances. Deer population density is often considered the main driver of browsing impacts on young trees, however, a range of other variables such as food availability also affect this relationship. In this study, we use browsing survey data from 135 research plots to explore patterns of roe deer (Capreolus capreolus) browsing pressure on woody plants in mountainous forests in central Europe. We fitted species-specific generalised linear mixed models for eight woody taxa, assessing the potential effects of understory characteristics, roe deer abundance and lying deadwood on browsing intensity. Our study reveals conspecific and associational effects for woody taxa that are intermediately browsed by roe deer. Selective browsing pressure was mediated by preferences of plants, in that, browsing of strongly preferred woody taxa as for example mountain ash (Sorbus aucuparia) and of least preferred woody taxa, for example Norway spruce (Picea abies) was not affected by the surrounding understory vegetation, while browsing pressure on intermediately browsed species like for example silver fir (Abies alba) was affected by understory characteristics. Contrary to our expectations, roe deer abundance was only positively associated with browsing pressure on silver fir and bilberry (Vaccinium myrtillus), while all other plants were unaffected by deer abundance. Finally, we did not find an influence of lying deadwood volume on the browsing pressure on any woody-plant species. Overall, our results indicate that patterns in browsing preference and intensity are species-specific processes and are partly affected by the surrounding understory vegetation. Current management strategies that aim to reduce browsing pressure through culling may be inefficient as they do not address other drivers of browsing pressure. However, managers also need to consider the characteristics of the local understory vegetation in addition to deer abundance and design species-specific plans to reduce browsing on woody plant taxa.

Interspecific Growth Reductions Caused by Wild Ungulates on Tree Seedlings and Their Implications for Temperate Quercus-Fagus Forests

Ungulate impacts on forest understory alter tree species composition, with cascading effects on forest functions and resilience against future climate conditions. Indeed, the ungulate browsing pressure on tree seedlings is species-specific and causes contrasted growth reductions that alter tree recruitment rates. Untangling the effects of browsing from the effects of the other factors driving regeneration success is required to guide the forest and ungulate management. In particular, Fagus sylvatica L. strongly dominates temperate Quercus-Fagus forests close to their climax, and it remains unclear if controlling ungulate populations can maintain tree species diversity in naturally regenerated forests. We addressed this question by monitoring 734 pairs of fenced and unfenced 6-m2 plots across a broad gradient of Cervus elaphus L. abundance in Belgian Quercus-Fagus forests managed by continuous cover forestry. Seedling height, density, and vegetation cover were monitored from 2016 to 2021. Species diversity and ecological affinity for light, temperature, and atmospheric humidity conditions were computed from these measures. With ungulates, the mean growth of Betula pendula Roth. and Sorbus aucuparia L. was negligible, whereas, without ungulates, their growth was higher than the growth of other species. With ungulates, the growth of Fagus sylvatica L. and Picea abies (L.) H. Karst was higher than other species. Quercus (Quercus petreae (Matt.) Liebl and Quercus robur L.) growth was the lowest in all conditions. Finally, Carpinus betulus L. was heavily browsed but still grew higher than its competitors with ungulates. Ungulate browsing can then severely affect seedling growth and likely reduce the diversity of future recruited trees. In the study area, browsing unfavored the regeneration of the species that are less shade tolerant, more-drought tolerant, and more-heat tolerant. It thus accelerates the natural succession and reduces forest resilience to heat and drought events. Such an observation was found valid over a wide study area encompassing contrasting levels of Cervus elaphus L. abundance. Combining further reductions of ungulate populations with foodscape improvement is likely required to maintain species diversity in these forests.

Restoring mixed forests through conversion of Norway spruce stands: effects of fencing and mechanical site preparation on performance of planted beech and natural tree regeneration

Conversion of Norway spruce (Picea abies) plantations to more diverse and resilient forest types is an important task for European forest managers in the face of climate change and increased focus on ecosystem services beyond timber production. However, there is a lack of knowledge on how to cost-effectively restore such forests. This study reports the influence of vicinity (distance) of forest type (mixed or spruce), fencing and mechanical site preparation (MSP) on the early performance of planted beech (Fagus sylvatica) seedlings and natural regeneration of other tree species following clear-cuts of Norway spruce in southern Sweden. After 6 years, we found clear effects of fencing and MSP, but not of vicinity of forest type. Fencing had a positive effect on height growth of Scots pine (Pinus sylvestris) but not on height of planted beech and naturally regenerated birch (Betula pendula, B. pubescens). There was a positive effect of MSP on survival and height growth of planted beech, and on the amount of natural regeneration of Scots pine. We conclude that establishment of beech forest is greatly accelerated by active regeneration approaches such as planting. In addition, the combination of planting beech, natural regeneration of other species, fencing and MSP is effective to promote the transition to mixed and diverse stands with both broadleaves and conifers. Fencing represented the highest cost among the treatments, and its cost-effectiveness depends on the local ungulate browsing pressure. In our study, fencing was critical to protect natural regeneration of Scots pine from browsing. Finally, natural regeneration of birch was abundant in our study and relatively unaffected by fencing and MSP treatments. With time, pre-commercial thinning of the naturally regenerated birch will be needed to maintain a diverse mixture of tree species.

Impact of white-tailed deer on northern white cedar recruitment beyond the browsing vulnerability zone: A broad-scale study

By selectively feeding on the new vegetation from forest regeneration, cervids influence the abundance of understory species. This can lead to recruitment failure for the consumed species. The impact of cervids on regeneration is generally studied at fine scales, making it difficult to generalize about the species distribution scale for co-occurring cervid and tree species. This is especially problematic for species such as northern white cedar (Thuja occidentalis L.), which is susceptible to browsing and has been declining throughout much of its range. In our study, we aimed to determine the effect of the spatio-temporal variations in white-tailed deer (Odocoileus virginianus Zimmerman) abundance on northern white cedar recruitment over a broad spatial scale. We combined distinct forest and wildlife monitoring databases for two Canadian provinces, Québec and New Brunswick. We also surveyed cedar regeneration and deer browsing pressure in a subset of plots located in Québec. In both provinces, we detected a negative effect of deer abundance and deeryard proximity on the recruitment of cedar saplings. In Québec, local deer browsing pressure also had a negative effect on cedar seedling abundance. As deer browsing pressure can vary considerably over a short distance, we recommend the inclusion of browsing indicators in national forest surveys to precisely assess the impact of browsing on regeneration and recruitment. This would allow managers to identify areas that are highly vulnerable to browsing and adapt forest management accordingly to reduce browsing-induced regeneration failures.

Escape Game: Responses of Northern White Cedar (Thuja occidentalis L.) to an Extreme Reduction in White-Tailed Deer (Odocoileus virginianus Zimmerman) Population

Northern white cedar (Thuja occidentalis L.) is a species of high ecological and economic value whose abundance has been declining since the pre-industrial period. It is an important element of white-tailed deer (Odocoileus virginianus Zimmerman) habitat, but its regeneration can be compromised by excessive browsing. This situation is especially critical in deeryards, where deer concentrate during winter. In 2018 and 2019, deer culling operations were carried out by the Ministry of Forests, Wildlife and Parks of Québec over 400 km2 in response to the occurrence of chronic wasting disease cases on a red deer farm. This operation offered an opportunity to look at how variations in deer pressure influence cedar regeneration and how cedar responds to a sudden reduction in browsing. We conducted regeneration surveys within and outside mapped deeryards both in the deer reduction zone and in a control zone. We performed dendrochronological analyses of cedar seedlings and saplings to quantify radial growth response to a reduction in browsing pressure. The results show that cedar basal area influences the abundance of small seedlings but that the effect of browsing becomes dominant for seedlings taller than 50 cm. Cedar growth responds to a reduction in browsing, but a two-year period was not sufficient to translate into changes in regeneration structure. The duration of the windows of opportunity at low browsing pressure required for cedar to reach a safe height remains to be determined. However, canopy openness had a significant influence on growth, suggesting that silvicultural measures could be taken to shorten the period of vulnerability to deer browsing.

Assessing the Heterogeneity and Conservation Status of the Natura 2000 Priority Forest Habitat Type Tilio–Acerion (9180*) Based on Field Mapping

Priority habitat types (HTs) within the Natura 2000 network are of the highest importance for conservation in Europe. However, they often occur in smaller areas and their conservation status is not well understood. One such HT is that of the Tilio–Acerion forests of slopes, screes and ravines (9180*). The Natura 2000 study site, Boč–Haloze–Donačka gora, in the Sub-Pannonian region of eastern Slovenia is characterized by a matrix of European beech forests and includes rather small, fragmented areas covered by Tilio–Acerion forests. The goal of this research was to examine the heterogeneity and conservation status of the selected HT through field mapping, which was performed in the summer of 2020. As the conservation of HT calls for a more detailed approach, we distinguished between the following four pre-defined habitat subtypes: (i) Acer pseudoplatanus-Ulmus glabra stands growing mostly in concave terrain, (ii) Fraxinus excelsior stands growing on slopes, (iii) Tilia sp. stands with thermophilous broadleaves occurring on ridges and slopes, (iv) Acer pseudoplatanus stands occurring on more acidic soils with an admixture of Castanea sativa. Field mapping information was complemented with the assessment of habitat subtype characteristics using remote sensing data. The results showed that habitat subtypes differed significantly in terms of area, tree species composition, forest stand characteristics, relief features and the various threats they experienced (e.g., fragmentation, tree mortality, ungulate browsing pressure). The differences between subtypes were also evident for LiDAR-derived environmental factors related to topography (i.e., terrain steepness and Topographic Position Index). This study provides a baseline for setting more realistic objectives for the conservation management of priority forest HTs. Due to the specificities of each individual habitat subtype, conservation activities should be targeted to the Natura 2000 habitat subtype level.