Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

The oak browsing index correlates linearly with roe deer density: a new indicator for deer management?

Increasing populations of large herbivores during the last decades have had a major impact on vegetation. While several studies have looked for quantifying this impact in terms of plant biomass, plant survival or financial costs, the potential benefit of using the response of the vegetation to changes in browsing pressure by large herbivores to monitor their populations has been poorly investigated. As getting accurate estimates of density in populations of large herbivores is problematic, the use of indicators measuring the intensity of browsing might offer reliable alternative to managers. From the intensive monitoring of a roe deer population subject to an experimental manipulation of density, we looked for assessing the response of oak to changes of roe deer population size. Using a simple browsing index calculated from field data over 10 years, we found that this oak browsing index linearly increased with increasing population size of roe deer. This suggests that such an oak browsing index might be a reliable “indicator of ecological change” for monitoring roe deer populations in oak forests with natural regeneration.     

Evaluation of potential deer browsing impact on sunflower (<Emphasis Type="Italic">Helianthus annus</Emphasis>)

Large herbivores are the key species for game management in the Central Europe. Analyses of factors affecting the feeding behaviour of herbivores and consequences of their browsing are therefore highly important both for farmers and for game managers as the protective measurements should be focused on the most threatened fields. The influence of fallow (Dama dama) and roe deer (Capreolus capreolus) browsing during vegetation period on sunflower production was studied. The experiment was carried out in 2006 on a field located near a forest complex. We marked out pairs of paired permanent plots and monitored deer impact regularly from the emergence of plants until the harvest. Herbivores damaged the sunflower intensively immediately after the emergence at the height of 1 to 2 cm above the ground. When the plants had reached approximately 15-40 cm in height, deer browsed the top shoots and as the plants grew higher the browsing was restricted to leaves or to substitute stems of the previously browsed plants. Sunflower ceased to attract big herbivores in flowering stage. Browsing significantly affected the yield of seeds. Approximately one half of the plants that had been damaged in early stage died as well as 12.5% of plants that had been damaged at second term; the rest produced substitute stems. Almost 33% of the substitute stems did not form disc florets; the second third of them formed disc florets with a diameter of 6-8 cm, and the last third set disc florets with a diameter of 8-10 cm.     

Ecological correlates of Himalayan musk deer Moschus leucogaster

Himalayan musk deer (Moschus leucogaster; hereafter musk deer) are endangered as a result of poaching and habitat loss. The species is nocturnal, crepuscular, and elusive, making direct observation of habitat use and behavior difficult. However, musk deer establish and repeatedly use the same latrines for defecation. To quantify musk deer habitat correlates, we used observational spatial data based on presence–absence of musk deer latrines, as well as a range of fine spatial‐scale ecological covariates. To determine presence–absence of musk deer, we exhaustively searched randomly selected forest trails using a 20‐m belt transect in different study sites within the Neshyang Valley in the Annapurna Conservation Area. In a subsequent way, study sites were classified as habitat or nonhabitat for musk deer. A total of 252 plots, 20 × 20 m, were systematically established every 100 m along 51 transects (each ~0.5 km long) laid out at different elevations to record a range of ecological habitat variables. We used mixed‐effect models and principal component analysis to characterize relationships between deer presence–absence data and habitat variables. We confirmed musk deer use latrines in forests located at higher elevations (3,200–4,200 m) throughout multiple seasons and years. Himalayan birch (Betula utilis) dominated forest, mixed Himalayan fir (Abies spectabilis), and birch forest were preferred over pure Himalayan fir and blue pine (Pinus wallichiana) forest. Greater crown cover and shrub diversity were associated with the presence of musk deer whereas tree height, diameter, and diversity were weakly correlated. Topographical attributes including aspect, elevation, distance to water source, and slope were also discriminated by musk deer. Over‐ and understory forest management can be used to protect forests likely to have musk deer as predicted by the models to ensure long‐term conservation of this rare deer.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

Simulated responses of moose populations to browsing‐induced changes in plant architecture and forage production

We and others have previously shown that browsing by large mammalian herbivores can alter the fractal dimension and other architectural properties of plant crowns. Using data from an experiment in northern Sweden which manipulated moose population densities from 0 to 50 moose per 1000 ha and measured architectural responses of birch, Betula pendula and B. pubescens, and Scots pine, Pinus sylvestris, we constructed a simulation model to examine how these changes in plant crown architecture affect moose population dynamics. Architectural changes in birch tree crowns caused forage availability to increase from low to moderate moose population densities, then decline at higher densities. In contrast, forage availability of pine decreased monotonically with increasing moose population density. The model, which incorporated equations fitted to these experimental results, predicted realistic moose population densities for northern Sweden. More importantly, the model also predicted that the quadratic responses of birch crowns to moose browsing results in moose population oscillations on high productivity sites because the moose population density overshoots the maximum birch forage availability then declines. Changes in the geometry of plant canopies caused by mammalian browsers and soil fertility may feed back on the population dynamics of the browsers themselves and even result in complex dynamics such as population oscillations.     

Associational refuge in practice: can existing vegetation facilitate woodland restoration?

Herbivores can dramatically diminish revegetation success, but associational refuge theory predicts that neighbouring plants could hinder browsing of planted seedlings. The key to strategic restoration using associational refuge is to define which patch variables are effective against the appropriate herbivores, at multiple scales, and to understand which stages of the foraging process these variables disrupt. Our study aimed to test the capacity of existing vegetation to act as associational refuge for planted seedlings by affecting search, detection and consumption decisions, and more generally influence herbivore foraging patterns. We conducted a field trial with free‐ranging, mammalian herbivores and nursery‐raised, native tree seedlings. We quantified seedling browsing damage over time in relation to a suite of existing patch variables at two spatial scales (100 m² and 4 m²). After two months, 78% of seedlings were browsed, suffering mean foliage loss of 90.5%. Focal seedlings were almost exclusively consumed by swamp wallabies Wallabia bicolor, an abundant generalist browser. Once a swamp wallaby investigated a seedling, the probability of consumption was high (86%). At the large scale, browsing of seedlings was delayed in patches with lower canopy cover and fewer browsed plant species. Seedlings in fern‐dominated patches escaped browsing for longer than those in grass‐dominated patches. At the small scale, browsing was delayed with higher cover of understorey vegetation. Associational refuge was provided by vegetation with characteristics, and at spatial scales, consistent with disrupted search and detection of focal seedlings by herbivores. Thus strategic placement of seedlings in existing vegetation – based on understanding which herbivore species is responsible and how it responds to vegetation – can take advantage of associational refuge during restoration. However, given rapid seedling detection by herbivores, associational refuge may be inadequate in the long‐term under high browsing pressure unless high absolute numbers of seedlings are planted among refuge.     

Exclusion of deer affects responses of birds to woodland regeneration in winter and summer

Using an exclosure experiment in managed woodland in eastern England, we examined species and guild responses to vegetation growth and its modification by deer herbivory, contrasting winter and the breeding season over 4 years. Species and guild responses, in terms of seasonal presence recorded by multiple point counts, were examined using generalized linear mixed models. Several guilds or migrant species responded positively to deer exclusion and none responded negatively. The shrub‐layer foraging guild was recorded less frequently in older and browsed vegetation, in both winter and spring. Exclusion of deer also increased the occurrence of ground‐foraging species in both seasons, although these species showed no strong response to vegetation age. The canopy‐foraging guild was unaffected by deer exclusion or vegetation age in either season. There was seasonal variation in the responses of some individual resident species, including a significantly lower occurrence of Eurasian Wren Troglodytes troglodytes and European Robin Erithacus rubecula in browsed vegetation in winter, but no effect of browsing on those species in spring. Ordinations of bird assemblage compositions also revealed seasonal differences in response to gradients of vegetation structure generated by canopy‐closure and exclusion of deer. Positive impacts of deer exclusion in winter are probably linked to reduced thermal cover and predator protection afforded by browsed vegetation, whereas species that responded positively in spring were also dependent on a dense understorey for nesting. The effects on birds of vegetation development and its modification by herbivores extend beyond breeding assemblages, with different mechanisms implicated and different species affected in winter.     

Regeneration of Salix arbuscula and Salix lapponum within a Large Mammal Exclosure: The Impacts of Microsite and Herbivory

Over‐grazing or browsing by large herbivores may result in the loss of individual plant species or entire plant communities. Restoration schemes often involve exclusion of large mammals, but the resulting changes in vegetation may alter other important ecological processes such as regeneration, via changes in microsite availability for seed germination or increases in populations of seedling predators. Working within a large fenced area from which large mammals were excluded, we experimentally tested the effects of microsite, small herbivores, and their interactions on post‐dispersal seed and early seedling mortality of one nationally scarce (Salix arbuscula) and one nationally rare (S. lapponum) species of montane willow. Seeds were sown in three different microsites: natural vegetation, mown vegetation (mimicking grazed sward), and bare ground. Small exclosures and slug pellets were used to examine the effects of small mammal and slug predation, respectively. Survival of seedlings was monitored during the summer following planting. The presence of bare ground, rather than the absence of herbivores, was of over‐riding importance for early seedling survival and establishment. Protecting seedlings from small mammals made no difference to the levels of survival; however, protecting seedlings from slugs (Arion spp.) resulted in approximately 45% of seedlings surviving until the end of the summer compared to only 30% when seedlings were available to slugs. Although excluding large herbivores may increase seed production of existing individuals, the impacts of changes to plant communities on processes such as regeneration need to be considered if restoration projects are to be fully successful.     

Establishment and Control of Hay-scented Fern: A Native Invasive Species

Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m² and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

Spatially biased reduction of browsing damage by sika deer through culling

Culling large herbivores can reduce browsing damage. Our objective was to verify the effect of culling by considering spatial changes in browsing damage to test the hypothesis that the benefits of spatially biased culling extend to the spatial scale of culling. Culling sika deer (Cervus nippon) in Kumamoto Prefecture, Japan, increased from 2009–2017, but browsing damage remained stable across the prefecture; regional damage trends differed among grid meshes (i.e., 5 km × 5 km). Meshes with browsing damage reduction received higher culling pressure and a decline in the deer population. Browsing damage reduction from culling was not uniform across regions and was biased by spatial bias in culling. This study highlights the importance of spatial scale in large herbivore management and evaluation of its effect.     

Positive interactions between herbivores and plant diversity shape forest regeneration

The effects of herbivores and diversity on plant communities have been studied separately but rarely in combination. We conducted two concurrent experiments over 3 years to examine how tree seedling diversity, density and herbivory affected forest regeneration. One experiment factorially manipulated plant diversity (one versus 15 species) and the presence/absence of deer (Odocoileus virginianus). We found that mixtures outperformed monocultures only in the presence of deer. Selective browsing on competitive dominants and associational protection from less palatable species appear responsible for this herbivore-driven diversity effect. The other experiment manipulated monospecific plant density and found little evidence for negative density dependence. Combined, these experiments suggest that the higher performance in mixture was owing to the acquisition of positive interspecific interactions rather than the loss of negative intraspecific interactions. Overall, we emphasize that realistic predictions about the consequences of changing biodiversity will require a deeper understanding of the interaction between plant diversity and higher trophic levels. If we had manipulated only plant diversity, we would have missed an important positive interaction across trophic levels: diverse seedling communities better resist herbivores, and herbivores help to maintain seedling diversity.     

Indirect interactions between browsers and seed predators affect the seed bank dynamics of a chaparral shrub

Interactions between herbivores and seed predators may have long-term consequences for plant populations that rely on persistent seed banks for recovery after unpredictable fires. We assessed the effects of browsing by deer and seed predation by rodents, ants and birds on the densities of seeds entering the seed bank of Ceanothus cuneatus var. rigidus, a maritime chaparral shrub in coastal California. Ceanothus produced many more seeds when protected from browsers in long-term experimental exclosures than did browsed plants, but the seed densities in the soil beneath browsed and unbrowsed Ceanothus were the same at the start of an intensive one-year study. The density of seeds in the soil initially increased in both treatments following summer seed drop: while densities returned to pre-drop levels within a few weeks under browsed plants, soil seed densities remained high for 5–8 months beneath unbrowsed plants. Rodent abundance (especially deer mice) was higher near unbrowsed plants than >30 m away, and rodents removed Ceanothus seeds from dishes in the experimental plots. At least in the short term, rodent density and rates of seed removal were inversely related to the intensity of browsing. Our data have management implications for maintaining viable Ceanothus populations by regulating the intensity of browsing and the timing, intensity and frequency of fires.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

Land abandonment and changes in snow cover period accelerate range expansions of sika deer

Ongoing climate change and land‐use change have the potential to substantially alter the distribution of large herbivores. This may result in drastic changes in ecosystems by changing plant–herbivore interactions. Here, we developed a model explaining sika deer persistence and colonization between 25 years in terms of neighborhood occupancy and habitat suitability. We used climatic, land‐use, and topographic variables to calculate the habitat suitability and evaluated the contributions of the variables to past range changes of sika deer. We used this model to predict the changes in the range of sika deer over the next 100 years under four scenario groups with the combination of land‐use change and climate change. Our results showed that both climate change and land‐use change had affected the range of sika deer in the past 25 years. Habitat suitability increased in northern or mountainous regions, which account for 71.6% of Japan, in line with a decrease in the snow cover period. Habitat suitability decreased in suburban areas, which account for 28.4% of Japan, corresponding to land‐use changes related to urbanization. In the next 100 years, the decrease in snow cover period and the increase in land abandonment were predicted to accelerate the range expansion of sika deer. Comparison of these two driving factors revealed that climate change will contribute more to range expansion, particularly from the 2070s onward. In scenarios that assumed the influence of both climate change and land‐use change, the total sika deer range increased by between +4.6% and +11.9% from the baseline scenario. Climate change and land‐use change will require additional efforts for future management of sika deer, particularly in the long term.     

The canary in the coal mine: Sprouts as a rapid indicator of browse impact in managed forests

Forest managers are frequently confronted with sustaining vegetation diversity and structure in landscapes experiencing high ungulate browsing pressure. Often, managers monitor browse damage and risk to plant communities using vegetation as indicators (i.e., phytoindicators). Although useful, the efficacy of traditional phytoindicators is sometimes hampered by limited distribution and abundance, variable browse susceptibility, and lagged responses. In contrast, sprouts possess traits which make them readily available and attractive to browsers, yet fairly resilient to tissue loss. Here, we experimentally evaluate whether hardwood tree stump sprouts are effective and sensitive phytoindicators of deer browse pressure. We measured sprout abundance and height in fenced and unfenced plots at 17 shelterwood harvested sites scattered across a 6500 km² region where deer densities varied by nearly an order of magnitude. We found browsing did not alter the proportion of stumps sprouting and sprout density; however, browse pressure varied among the four most abundant species. Acer rubrum and Acer saccharum were heavily browsed, although browse pressure on A. saccharum decreased in areas with greater canopy openness. Fagus grandifolia and Prunus serotina were less preferred. Differences in palatability altered size hierarchies. Averaged across all species, browsing reduced sprout height by 39%, relative to protected sprouts. Under ambient browsing, P. serotina was 60–100% taller than other species and significantly taller than A. saccharum and F. grandifolia. However, within fences A. saccharum and A. rubrum doubled in size, relative to browsed individuals, and were as tall as P. serotina. Deer impact on sprout height within unfenced forest stands was negatively correlated with estimated deer densities (R² = 0.46). Thus, we suggest sprout surveys can provide a measure of impact across much larger areas. Our results demonstrate that sprouts, particularly those of Acer species, offer an abundant, easily measured, and reliable indicator of browse pressure. Moreover, browse impacts on sprouts emerged before impacts were detected on seedling abundance, height, or biomass. We argue sprouts can warn of imminent browse risk to seedlings (and perhaps non-woody vegetation) and thereby allow managers to take actions to mitigate or avert losses to the regenerating seedling cohort.     

Disentangling the causes of damage variation by deer browsing on young Thuja plicata

Long‐lived trees experience different levels of damage due to mammalian herbivores. To untangle the mechanisms that underlie this variation, we combined chemical with dendrochronological analyses to study variation in browsing on Western redcedars (Thuja plicata) on Haida Gwaii (British Columbia, Canada). Since the last glaciation, Haida Gwaii forests had lacked large herbivore browser until Sitka black‐tailed deer (Odocoileus hemionus sitkensis) were introduced at the beginning of the 20th century. Dendrochronology yielded information on radial growth and plant annual responses to environmental stresses including herbivory. Secondary metabolite content and plant nutritional quality provided insights into proximate causes of food choices made by herbivores. We sampled lightly‐ and heavily‐browsed young trees at four sites: three clear‐cut sites with high browsing pressure and one old‐growth forest site where browsing pressure had, until recently, been lower. Heavily‐browsed young trees had lower concentrations of secondary metabolites and were of lower nutritive value than lightly‐browsed trees at all sites. Under high browsing pressure, tree growth patterns suggested that all young trees were initially severely browsed until some trees, currently scored as lightly‐browsed, started to escape deer. At the old‐growth site, both lightly‐ and heavily‐browsed trees tended to have lower overall average secondary metabolite concentrations than those of all other sites, a trend possibly related to greater canopy closure. Lightly‐browsed trees were older than heavily‐browsed ones which resulted, during the period of lower browsing pressure, in higher growth rate and a same pattern of change in growth from one year to the next year. This suggests that, under low browsing pressure, selection of young trees related to chemical defense was weak and that growth differences due to other factors than browsing could be expressed. Under strong browsing pressure, however, all young trees had equally low growth rates until trees with better genetic potential to produce effective defenses were able to escape deer. This suggests that selection by deer could occur on a long‐lived tree.     

Herbivory on planted oak seedlings across a habitat edge created by timber harvest

Edge habitats create environmental gradients that affect plant community composition and herbivore behavior. Silvicultural disturbance creates edge habitat with direct (via changes in light) and indirect (via changes in herbivore behavior) consequences for the growth and survival of tree seedlings, and thus, the composition of the future forest stands. Herbivores, particularly ungulates, can be a major limiting factor in oak regeneration, and silvicultural disturbance may alter the abundance or behavior of herbivores following harvest. We measured the severity of herbivory on experimentally planted white (Quercus alba) and black oak (Quercus velutina) seedlings by white-tailed deer (Odocoileus virginianus) and eastern cottontail rabbits (Sylvilagus floridanus), as well as foliar damage from insects, across gradients created by clearcuts in a deciduous forest in Indiana, USA. Overall browse pressure on oaks was low in our study. Nonetheless, spatial variation in herbivory depended on herbivore taxa; herbivory by rabbits was highest inside harvest openings, whereas foliar damage by insects peaked in the forest. Intensity of deer herbivory was constant across the edge. In addition, we observed indirect interactions among herbivore species mediated by a seedling’s browsing history. Herbivore damage by deer was positively related to past browsing by rabbits, and foliar damage from insects was positively related to past browsing by both deer and rabbits. Increasing woody plant competition reduced herbivory on seedlings by both deer and rabbits. Given the lack of spatial variability in deer herbivory and low overall herbivory by rabbits, we suspect that interactions between timber harvesting and herbivory did not have a strong impact on oak seedlings at our study sites.     

White-tailed deer (Odocoileus virginianus) fecal pellet decomposition is accelerated by the invasive earthworm Lumbricus terrestris

Exotic European earthworms have expanded into worm-free forests of the United States. Concurrently, populations of the white-tailed deer, Odocoileus virginianus, have also increased. During winter, deer use hemlock stands for cover while browsing elsewhere, creating a net organic matter flux into these stands. Deer fecal pellets can provide annual inputs of 48.1 kg C, 1.4 kg N, and 1.3 kg Ca per hectare. We tested the hypothesis that these pellets were readily consumed by invading earthworms. The litter-feeding anecic earthworm Lumbricus terrestris redistributed fecal pellets and accelerated mass and nutrient loss rates. These losses are likely due to the combination of enhanced fragmentation and decomposition as earthworms drag pellets into their burrows for consumption. This nutrient subsidy may be an important source of high quality “litter” input to hemlock stands, which may in turn facilitate the invasion of these stands by earthworms under high deer densities.