A Natural Experiment on the Impact of Overabundant Deer on Forest Invertebrates

Abstract:  In large parts of North America and Europe, deer overabundance threatens forest plant diversity. Few researchers have examined its effects on invertebrate assemblages. In a natural experiment on Haida Gwaii (British Columbia, Canada), where Sitka black-tailed deer ( Odocoileus hemionus sitkensis ) were introduced, we compared islands with no deer, with deer for fewer than 20 years, and with deer for more than 50 years. We sampled invertebrates in three habitat categories: forest edge vegetation below the browse line, forest interior vegetation below the browse line, and forest interior litter. In forest edge vegetation, invertebrate abundance and species density decreased with increasing length of browsing history. In forest interior vegetation, decrease was significant only on islands with more than 50 years of browsing. Insect abundance in the vegetation decreased eightfold and species density sixfold on islands browsed for more than 50 years compared with islands without deer. Primary consumers were most affected. Invertebrates from the litter showed little or no variation related to browsing history. We attributed the difference between vegetation-dwelling and litter-dwelling invertebrates to differences in the effect of browsing on their habitat. In the layer below the browse line deer progressively removed the habitat. The extent of litter habitat was not affected, but its quality changed. We recommend more attention be given to the effect of overabundant ungulates on forest invertebrate conservation with a focus on edge and understory vegetation in addition to litter habitat.     

Response of a Reptile Guild to Forest Harvesting

Abstract:  Despite the growing concern over reptile population declines, the effects of modern industrial silviculture on reptiles have been understudied, particularly for diminutive and often overlooked species such as small-bodied snakes. We created 4 replicated forest-management landscapes to determine the response of small snakes to forest harvesting in the Coastal Plain of the southeastern United States. We divided the replicated landscapes into 4 treatments that represented a range of disturbed habitats: clearcut with coarse woody debris removed; clearcut with coarse woody debris retained; thinned pine stand; and control (unharvested second-growth planted pines). Canopy cover and ground litter were significantly reduced in clearcuts, intermediate in thinned forests, and highest in unharvested controls. Bare soil, maximum air temperatures, and understory vegetation all increased with increasing habitat disturbance. Concomitantly, we observed significantly reduced relative abundance of all 6 study species (scarletsnake [Cemophora coccinea] , ring-neck snake [Diadophis punctatus] , scarlet kingsnake [Lampropeltis triangulum] , red-bellied snake [Storeria occipitomaculata] , southeastern crowned snake [Tantilla coronata] , and smooth earthsnake [Virginia valeriae] ) in clearcuts compared with unharvested or thinned pine stands. In contrast, the greatest relative snake abundance occurred in thinned forest stands. Our results demonstrate that at least one form of forest harvesting is compatible with maintaining snake populations. Our results also highlight the importance of open-canopy structure and ground litter to small snakes in southeastern forests and the negative consequences of forest clearcutting for small snakes.     

Using forest inventory data to assess fisher resting habitat suitability in California.

The fisher (Martes pennanti) is a forest-dwelling carnivore whose current distribution and association with late-seral forest conditions make it vulnerable to stand-altering human activities or natural disturbances. Fishers select a variety of structures for daily resting bouts. These habitat elements, together with foraging and reproductive (denning) habitat, constitute the habitat requirements of fishers. We develop a model capable of predicting the suitability of fisher resting habitat using standard forest vegetation inventory data. The inventory data were derived from Forest Inventory and Analysis (FIA), a nationwide probability-based sample used to estimate forest characteristics. We developed the model by comparing vegetation and topographic data at 75 randomly selected fisher resting structures in the southern Sierra Nevada with 232 forest inventory plots. We collected vegetation data at fisher resting locations using the FIA vegetation sampling protocol and centering the 1-ha FIA plot on the resting structure. To distinguish used and available inventory plots, we used nonparametric logistic regression to evaluate a set of a priori biological models. The top model represented a dominant portion of the Akaike weights (0.87), explained 31.5% of the deviance, and included the following variables: average canopy closure, basal area of trees <51 cm diameter breast height (dbh), average hardwood dbh, maximum tree dbh, percentage slope, and the dbh of the largest conifer snag. Our use of routinely collected forest inventory data allows the assessment and monitoring of change in fisher resting habitat suitability over large regions with no additional sampling effort. Although models were constrained to include only variables available from the list of those measured using the FIA protocol, we did not find this to be a shortcoming. The model makes it possible to compare average resting habitat suitability values before and after forest management treatments, among administrative units, across regions and over time. Considering hundreds of plot estimates as a sample of habitat conditions over large spatial scales can bring a broad perspective, at high resolution, and efficiency to the assessment and monitoring of wildlife habitat.     

Forests Too Deer: Edge Effects in Northern Wisconsin

Abstract: Browsing by white-tailed deer (Odocoileus virginianus) can profoundly affect the abundance and population structure of several woody and herbaceous plant species. Enclosure studies and population surveys reveal that past and current deer densities as low as 4 deer/km² may prevent regeneration of the once common woody species, Canada yew (Taxus canadensis), eastern hemlock (Tsuja canadensis), and white cedar Puja occidentalis), as well as several herbaceous species. Prior to European settlement, forests in northern Wisconsin contained relatively sparse deer populations (<4/km²), but extensive timber cutting in the late nineteenth century boosted deer populations. Continued habitat fragmentation resulting from scattered timber harvests and the creation of "wildlife openings" to improve deer forage maintain these high densities throughout much of the Northeast.
Because deer wander widely, the effects of high deer densities penetrate deeply into remaining stands of old and mature forest, greatly modifying their composition Thus, abundant early successional and "edge" habitat, and the high deer densities they engender, represent significant external threats to these plant communities. We hypothesize that establishing large (200–400 km²) continuous areas of maturing forest, especially in conjunction with increased hunting, could reduce local deer densities and so provide a simple and inexpensive method for retaining species sensitive to the deleterious effects of browsing.     

Plethodontid Salamander Response to Silvicultural Practices in Missouri Ozark Forests

Abstract: There is little information on the effects of tree harvest on salamander populations in the midwestern United States. We present data on plethodontid salamander densities in replicated stands of three forest age classes in the southeastern Ozarks of Missouri. Forest age classes consisted of regeneration-cut sites <5 years old, second-growth sites 70–80 years old, and old-growth sites> 120 years old. Salamander abundance on 21, 144-m² plots was determined by area- and time-constrained searches. We also compared age-class habitat characteristics, including downed woody debris, canopy cover, ground area cover, herbaceous vegetation, and woody vegetation. Salamander density was lowest in newly regenerated forests and highest in forests> 120 years old. Comparisons of recently regenerated forests with mature forests> 70 years old indicated that terrestrial salamanders were reduced to very low numbers when mature forests had been intensively harvested. This reduction may result from a decrease in microhabitat availability. Forest age-class comparisons further indicated that salamander abundance slowly increased over time after forests had regenerated. Management decisions that take into account plethodontid salamander abundance and their response to forest structural diversity are important components in sustaining ecosystem integrity while maximizing economic yield.     

Cumulative ecological and socioeconomic effects of forest policies in coastal Oregon.

Forest biodiversity policies in multi-ownership landscapes are typically developed in an uncoordinated fashion with little consideration of their interactions or possible unintended cumulative effects. We conducted an assessment of some of the ecological and socioeconomic effects of recently enacted forest management policies in the 2.3-million-ha Coast Range Physiographic Province of Oregon. This mountainous area of conifer and hardwood forests includes a mosaic of landowners with a wide range of goals, from wilderness protection to high-yield timber production. We projected forest changes over 100 years in response to logging and development using models that integrate land use change and forest stand and landscape processes. We then assessed responses to those management activities using GIS models of stand structure and composition, landscape structure, habitat models for focal terrestrial and aquatic species, timber production, employment, and willingness to pay for biodiversity protection. Many of the potential outcomes of recently enacted policies are consistent with intended goals. For example, we project the area of structurally diverse older conifer forest and habitat for late successional wildlife species to strongly increase. 'Other outcomes might not be consistent with current policies: for example, hardwoods and vegetation diversity strongly decline within and across owners. Some elements of biodiversity, including streams with high potential habitat for coho salmon (Oncorhynchus kisutch) and sites of potential oak woodland, occur predominately outside federal lands and thus were not affected by the strongest biodiversity policies. Except for federal lands, biodiversity policies were not generally characterized in sufficient detail to provide clear benchmarks against which to measure the progress or success. We conclude that land management institutions and policies are not well configured to deal effectively with ecological issues that span broad spatial and temporal scales and that alternative policies could be constructed that more effectively provide for a mix of forest values from this region.     

Spatio-temporal modelling of ground vegetation development in mountain spruce forests

More complex models of forest ecosystems are required to understand how land-cover changes can impact vegetation dynamics and spatial pattern. In order to document spatio-temporal modelling abilities, the observations conducted in the declined climax mountain Norway spruce forest during the recovery period (1995-2006) are used for simulation and spatial analysis in the GIS environment. The developed spatio-temporal model is used for simulation of forest vegetation dynamics in a mountain spruce forest in the framework of regeneration processes after stress from air pollution. In order to explore the spatial and temporal phenomena of regeneration processes, the spatio-temporal model is based on a large set of ordinary differential equations that solve dynamic processes in sets of microsites arranged in grids for each ground vegetation species and each age group of Norway spruce seedlings. The spatial extent of the explored site is composed of a set of 50 × 50 microsites. Each microsite is represented by a square with dimensions of 1 m × 1 m. The presented simulation studies are mainly focused on seedlings from the seed year 1992, in order to explore the longest monitored time series of survival. It is based on exponential growth models that are related to the environmental conditions for each microsite. The canopy gaps based on estimates of the local crown projected area, the soil type layer, and the dominant grass density are used to provide case simulation studies. The first case study simulates the influence of microsite positions in relation to the local tree crown projections on the survival of spruce seedlings. It is assumed that the density of the trees is the main factor that determines the light and heat supply to the ground level of the Norway spruce seedlings. The second case study extends the previous study to include terms that determine the growth ratio in dependence on the crown projection area. The third case study provides further extensions in order to simulate growth ratio relations to the local soil type. The fourth case study demonstrates the local influence of the dominant grasses, such as Avenella flexuosa and Calamagrostis villosa, on the natural regeneration of Norway spruce. Starting from the conditions at the sites before the recovery period, the case simulation studies are able to project the short-term succession for a regeneration decade and the approximate long-term development. In addition to the standard simulation procedures based on solution of ordinary differential equations, spatio-temporal modelling in the GIS environment is able to provide spatial data management, analysis and visualization of the data.     

The effect of deer management on the abundance of Ixodes ricinus in Scotland

The management of wildlife hosts for controlling parasites and disease has a history of mixed success. Deer can be important hosts for ticks, such as Ixodes ricinus, which is the primary vector of disease-causing zoonotic pathogens in Europe. Deer are generally managed by culling and fencing for forestry protection, habitat conservation, and commercial hunting, and in this study we test whether these deer management methods can be useful for controlling ticks, with implications for tick-borne pathogens. At different spatial scales and habitats we tested the hypotheses that tick abundance is reduced by (1) culling deer and (2) deer exclusion using fencing. We compared abundance indices of hosts and questing I. ricinus nymphs using a combination of small-scale fencing experiments on moorland, a large-scale natural experiment of fenced and unfenced pairs of forests, and cross-sectional surveys of forest and moorland areas with varying deer densities. As predicted, areas with fewer deer had fewer ticks, and fenced exclosures had dramatically fewer ticks in both large-scale forest and small-scale moorland plots. Fencing and reducing deer density were also associated with higher ground vegetation. The implications of these results on other hosts, pathogen prevalence, and disease risk are discussed. This study provides evidence of how traditional management methods of a keystone species can reduce a generalist parasite, with implications for disease risk mitigation.     

Potential effects of forest policies on terrestrial biodiversity in a multi-ownership province.

We used spatial simulation models to evaluate how current and two alternative policies might affect potential biodiversity over 100 years in the Coast Ranges Physiographic Province of Oregon. This 2.3-million-ha province is characterized by a diversity of public and private forest owners, and a wide range of forest policy and management objectives. We evaluated habitat availability for seven focal species representing different life histories. We also examined how policies affected old-growth stand structure, age distributions relative to the historical range of variability, and landscape patterns of forest types. Under the current policy scenario, the area of habitat for old-growth forest structure and associated species increased over time, the habitat for some early-successional associates remained stable, and the area of hardwood vegetation and diverse early-successional stages declined. The province is projected to move toward but not reach the historical range of variation of forest age classes that may have occurred under the wildfire regimes of the pre-Euroamerican settlement period. Ownership explained much of the pattern of biodiversity in the province, and under the current policy scenario, its effect increased over time as the landscape diverged into highly contrasting forest structures and ages. Patch type diversity declined slightly overall but declined strongly within ownerships. Most of the modeled change in biodiversity over time resulted from policies on public forest lands that were intended to increase the area of late-successional forests and species. One of the alternative policies, increased retention of wildlife trees on private lands, reduced the contrast between ownerships and increased habitat availability over time for both early- and late-successional species. Analysis of another alternative, stopping thinning of plantations on federal lands, indicated that current thinning regimes improve habitat for the Olive-sided Flycatcher, but the no-thinning alternative had no effect on the habitat scores for the late-successional species in the 100-year simulation. A comparison of indicators of biological diversity suggests that using focal species and forest structural measures can provide complementary information on biodiversity. The multi-ownership perspective provided a more complete synthesis of province-wide biodiversity patterns than assessments based on single ownerships.     

The importance of spatial autocorrelation,extent and resolution in predicting forest bird occurrence

Concerns about declines in forest biodiversity underscore the need for accurate estimates of the distribution and abundance of organisms at large scales and at resolutions that are fine enough to be appropriate for management. This paper addresses three major objectives: (i) to determine whether the resolution of typical air photo-derived forest inventory is sufficient for the accurate prediction of site occupancy by forest birds. We compared prediction success of habitat models using air photo variables to models with variables derived from finer resolution, ground-sampled vegetation plots. (ii) To test whether incorporating spatial autocorrelation into habitat models via autologistic regression increases prediction success. (iii) To determine whether landscape structure is an important factor in predicting bird distribution in forest-dominated landscapes. Models were tested locally (Greater Fundy Ecosystem [GFE]) using cross-validation, and regionally using an independent data set from an area located ca. 250 km to the northwest (Riley Brook [RB]). We found significant positive spatial autocorrelation in the residuals of at least one habitat model for 76% (16/21) of species examined. In these cases, the logistic regression assumption of spatially independent errors was violated. Logistic models that ignored spatial autocorrelation tended to overestimate habitat effects. Though overall prediction success was higher for autologistic models than logistic models in the GFE, the difference was only significantly improved for one species. Further, the inclusion of spatial covariates did little to improve model performance in the geographically discrete study area. For 62% (13/21) of species examined, landscape variables were significant predictors of forest bird occurrence even after statistically controlling for stand-level variability. However, broad spatial extents explained less variation than local factors. In the GFE, 76% (16/21) of air photo and 81% (17/21) of ground plot models were accurate enough to be of practical utility (AUC > 0.7). When applied to RB, both model types performed effectively for 55% (11/20) of the species examined. We did not detect an overall difference in prediction success between air photo and ground plot models in either study area. We conclude that air photo data are as effective as fine resolution vegetation data for predicting site occupancy for the majority of species in this study. These models will be of use to forest managers who are interested in mapping species distributions under various timber harvest scenarios, and to protected areas planners attempting to optimize reserve function.     

Habitat and population modelling of roe deer using an interactive geographic information system

Management of German roe deer (Capreolus capreolus) populations is a challenge for wildlife managers and foresters because population densities are difficult to estimate in forests and forest regeneration can be negatively affected when roe deer density is high. We describe a model to determine deer population densities compatible with forest management goals, and to assess harvest rates necessary to maintain desired deer densities. A geographic information system (GIS) was used to model wildlife habitat and population dynamics over time. Our model interactively incorporates knowledge of field biologists and foresters via a graphical user interface (GUI). Calibration of the model with deer damage maps allowed us to evaluate density dependence of a roe deer population. Incorporation of local knowledge into temporally dynamic and spatial models increases understanding of population dynamics and improves wildlife management.     

Ecological consequences of forest elephant declines for Afrotropical forests

Poaching is rapidly extirpating African forest elephants (Loxodonta cyclotis) from most of their historical range, leaving vast areas of elephant‐free tropical forest. Elephants are ecological engineers that create and maintain forest habitat; thus, their loss will have large consequences for the composition and structure of Afrotropical forests. Through a comprehensive literature review, we evaluated the roles of forest elephants in seed dispersal, nutrient recycling, and herbivory and physical damage to predict the cascading ecological effects of their population declines. Loss of seed dispersal by elephants will favor tree species dispersed abiotically and by smaller dispersal agents, and tree species composition will depend on the downstream effects of changes in elephant nutrient cycling and browsing. Loss of trampling and herbivory of seedlings and saplings will result in high tree density with release from browsing pressures. Diminished seed dispersal by elephants and high stem density are likely to reduce the recruitment of large trees and thus increase homogeneity of forest structure and decrease carbon stocks. The loss of ecological services by forest elephants likely means Central African forests will be more like Neotropical forests, from which megafauna were extirpated thousands of years ago. Without intervention, as much as 96% of Central African forests will have modified species composition and structure as elephants are compressed into remaining protected areas. Stopping elephant poaching is an urgent first step to mitigating these effects, but long‐term conservation will require land‐use planning that incorporates elephant habitat into forested landscapes that are being rapidly transformed by industrial agriculture and logging.     

川西高山森林生态系统林下生物量及其随林窗的变化特征

作为森林生态系统的重要组成部分,林下植被及其残体的分布受到林冠层的影响,但迄今有关林窗对林下植被和残体生物量的影响尚无研究报道.于2013 年8 月2 日至20 日,以海拔3 600 m 的川西岷江冷杉原始林林下植被为研究对象,根据区域内的坡向和林分组成等因素设置3 个100 m×100 m 的典型样地,调查其生物量及其随林窗的变化特征.在每个样地内选择3 个大林窗,在林窗、林缘和林下分别设置3 个20 m×20 m 的样方,调查粗木质残体长度或高度、大小头直径、枯立木记录胸径、腐烂等级等;在林窗、林缘和林下分别设置3 个5 m×5 m 的样方,采用“收获法”收集样方内直径在2.5-10 cm之间的细木质残体和灌木生物量;在林窗、林缘和林下分别设置3 个1 m×1 m 的样方来调查凋落物储量和草本生物量;在1m×1 m 的样方内随机选择1 个20 cm×20 cm 的小样方来调查地被植物生物量.结果表明,（1）川西高山森林生态系统总生物量为72.75 t·hm^-2,其中林下生物量为67.92 t·hm^-2,占生态系统生物量的95.17%.活体植被以灌木为主,其生物量为9.81t·hm^-2;残体部分以粗木质残体为主,其储量为53.00 t·hm^-2;（2）林窗对灌木、草本、地被植物的影响各不相同,且不同物种的灌木生物量表现出不同的分布规律;草本生物量表现出明显的“边缘效应”,在林缘显著高于林下;林窗和林缘的地被植物生物量相对较低;（3）粗木质残体储量从林下到林窗呈现减小的趋势,但总体储量仍然较大,林窗和林缘的细木质残体储量高于林下.这些结果为认识高山森林生态系统林下生物量及其格局,以及林窗在森林生态系统的重要作用提供了基础理论依据.     

Community occupancy responses of small mammals to restoration treatments in ponderosa pine forests, northern Arizona, USA

In western North American conifer forests, wildfires are increasing in frequency and severity due to heavy fuel loads that have accumulated after a century of fire suppression. Forest restoration treatments (e.g., thinning and/or burning) are being designed and implemented at large spatial and temporal scales in an effort to reduce fire risk and restore forest structure and function. In ponderosa pine (Pinus ponderosa) forests, predominantly open forest structure and a frequent, low-severity fire regime constituted the evolutionary environment for wildlife that persisted for thousands of years. Small mammals are important in forest ecosystems as prey and in affecting primary production and decomposition. During 2006-2009, we trapped eight species of small mammals at 294 sites in northern Arizona and used occupancy modeling to determine community responses to thinning and habitat features. The most important covariates in predicting small mammal occupancy were understory vegetation cover, large snags, and treatment. Our analysis identified two generalist species found at relatively high occupancy rates across all sites, four open-forest species that responded positively to treatment, and two dense-forest species that responded negatively to treatment unless specific habitat features were retained. Our results indicate that all eight small mammal species can benefit from restoration treatments, particularly if aspects of their evolutionary environment (e.g., large trees, snags, woody debris) are restored. The occupancy modeling approach we used resulted in precise species-level estimates of occupancy in response to habitat attributes for a greater number of small mammal species than in other comparable studies. We recommend our approach for other studies faced with high variability and broad spatial and temporal scales in assessing impacts of treatments or habitat alteration on wildlife species. Moreover, since forest planning efforts are increasingly focusing on progressively larger treatment implementation, better and more efficiently obtained ecological information is needed to inform these efforts.     

Spatial occurrence of a habitat-tracking saproxylic beetle inhabiting a managed forest landscape.

Because of the dynamic nature of many managed habitats, proper evaluation of conservation efforts calls for models that take into account both spatial and temporal habitat dynamics. We develop a metapopulation model for successional-type systems, in which habitat quality changes over time in a predictable fashion. The occupancy and recruitment of the predatory saproxylic (dependent on dead wood) beetle Harminius undulatus was studied in a managed boreal forest landscape, covering 24,449 ha, in central Sweden. In a first step, we analyzed the beetle's occupancy pattern in relation to stand characteristics, and the amounts of present and past habitat in the surrounding landscape. Managed forest is suitable habitat when > or =60 years old, and immediately after cutting, but not between the ages of 10 and 60 years. The observed occupancy of H. undulatus was positively correlated with the stand's age as habitat. We used a metapopulation model to predict the current probability of occurrence in each forest stand, given the spatiotemporal distribution of suitable forest stands during the last 50 years. Metapopulation parameters were estimated by matching predicted spatial distributions with observed spatial distributions. The model predicted observed spatial distributions better than a similar model that assumed constant habitat quality of each forest stand. Thus, metapopulation models for successional-type systems, such as dead wood dependent organisms in managed forest landscapes, should include habitat dynamics. An estimated 82% of the landscape-wide recruitment took place in managed stands, which covered 87% of the forest area, in comparison with 18% in unmanaged stands, which covered 13% of the forest area. Among the managed stand types, > or =60-year-old stands and 3-7-year-old clear-cuttings contributed to 79% of the total recruitment while 8-59-year-old stands only contributed 3%. The results suggest the following guidelines to improve conditions for H. undulatus and other species with similar habitat requirements: (1) the proportion of the landscape constituted by younger stands should not be allowed to grow too large, (2) the rotation period of managed stands should not be allowed to be too short, and (3) dead wood should be retained and created at final cutting.     

Effects of Human Subsistence Activities on Forest Birds in Northern Kenya

Abstract: Indigenous tribes and conservation biologists may have common goals and may be able to collaborate on the maintenance of biodiversity, but few researchers have evaluated the impacts and potential benefits of human subsistence activities. I studied the effects of subsistence activities (primarily wood collection) of nomadic pastoralists in 3 Afromontane forests of northern Kenya. In surveys of 404, 25‐m‐radius plots, I recorded vegetation structure and composition of the forest bird community. Plots with higher levels of human activity had significantly different vegetation structure, with more‐open canopies, more grass, and fewer tree stems. Nectarivores (abundance +231%) and aerial insectivores (+66%) were more abundant in plots with more‐intense wood collecting than in plots with less human activity, whereas abundance of forest specialists (?28%) decreased in plots with more‐intense human activity. Abundance of 58% of the bird species either increased or decreased significantly in plots with more‐intense human activity. Generally, the number of individuals of forest specialists decreased (6 of 7 species showed significant responses) and the number of individuals of edge and nonforest species increased with increasing human activity. Canonical correspondence analysis showed that an intensification of human activities would favor nectarivores, aerial insectivores, granivores, and omnivores and would negatively affect large‐sized, ground‐foraging species and arboreal frugivores. Subsistence human activities favored the invasion of forest by edge species at the expense of forest specialists; thus, further intensification of forest exploitation by local peoples is not recommended. At the same time, however, subsistence activities in northern Kenya forests appeared to increase the structural diversity of the vegetation and provided suitable habitat for part (but not all) of the forest avifauna, which suggests that subsistence human activities may have a role in the maintenance of bird diversity.     

Modelling forest management within a global vegetation model—Part 1: Model structure and general behaviour

This article describes a new forest management module (FMM) that explicitly simulates forest stand growth and management within a process-based global vegetation model (GVM) called ORCHIDEE. The net primary productivity simulated by ORCHIDEE is used as an input to the FMM. The FMM then calculates stand and management characteristics such as stand density, tree size distribution, tree growth, the timing and intensity of thinnings and clear-cuts, wood extraction and litter generated after thinning. Some of these variables are then fed back to ORCHIDEE. These computations are made possible with a distribution-based modelling of individual tree size. The model derives natural mortality from the relative density index (rdi), a competition index based on tree size and stand density. Based on the common forestry management principle of avoiding natural mortality, a set of rules is defined to calculate the recurrent intensity and frequency of forestry operations during the stand lifetime. The new-coupled model is called ORCHIDEE-FM (forest management).The general behaviour of ORCHIDEE-FM is analysed for a broadleaf forest in north-eastern France. Flux simulation throughout a forest rotation compare well with the literature values, both in absolute values and dynamics.Results from ORCHIDEE-FM highlight the impact of forest management on ecosystem C-cycling, both in terms of carbon fluxes and stocks. In particular, the average net ecosystem productivity (NEP) of 225 gC m⁻² year⁻¹ is close to the biome average of 311 gC m⁻² year⁻¹. The NEP of the “unmanaged” case is 40% lower, leading us to conclude that management explains 40% of the cumulated carbon sink over 150 years. A sensitivity analysis reveals 4 major avenues for improvement: a better determination of initial conditions, an improved allocation scheme to explain age-related decline in productivity, and an increased specificity of both the self-thinning curve and the biomass-diameter allometry.     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.     

Managing the Abundance and Diversity of Breeding Bird Populations through Manipulation of Deer Populations

Abstract: Deer densities in forests of eastern North America are thought to have significant effects on the abundance and diversity of forest birds through the role deer play in structuring forest understories. We tested the ability of deer to affect forest bird populations by monitoring the density and diversity of vegetation and birds for 9 years at eight 4-ha sites in northern Virginia, four of which were fenced to exclude deer. Both the density and diversity of understory woody plants increased following deer exclosure. The numerical response of the shrubs to deer exclosure was significantly predicted by the soil quality (ratio of organic carbon to nitrogen) at the sites. Bird populations as a whole increased following exclosure of deer, particularly for ground and intermediate canopy species. The diversity of birds did not increase significantly following exclosure of deer, however, primarily because of replacement of species as understory vegetation proceeded through successional processes. Changes in understory vegetation accounted for most of the variability seen in the abundance and diversity of bird populations. Populations of deer in protected areas are capable of causing significant shifts in the composition and abundance of bird communities. These shifts can be reversed by increasing the density and diversity of understory vegetation, which can be brought about by reducing deer density.