Tree shelters facilitate brown oak seedling survival and establishment in a grazing‐dominant forest of Bhutan,Eastern Himalaya

Brown oak (Quercus semecarpifolia) forest is essential for ecological and socioeconomic functions, mainly grazing in the Himalayas. The tree has failed to regenerate naturally and is a threatened species. Restoration of brown oaks is crucial to ensure sustainability while maintaining livestock grazing in these habitats. Achieving this requires cost‐effective restoration techniques that are practicable and sympathetic to the multiple uses of the forest. We assessed the combined effect of grazing (control) and three tree shelters (Protex tubes, mesh wires, and wooden frames) on the field performance of oak seedlings in a forest with heavy grazing pressure. Seedling survival and morphological indicators, including seedling height, collar diameter, sturdiness quotient (SQ), and leaf mass per area (LMA) indices, were measured. More than 90% of control seedlings without protective shelters suffered severe browsing and demonstrated significantly lower survival rates compared to tree shelter seedlings, indicating that grazing was the primary factor governing regeneration success. Seedling survival in tree shelters was three times higher, while the height increase was two times higher than the control. Additionally, locally made mesh wire and wooden tree shelters were more effective than Protex and control in producing quality seedlings reflected by the SQ and LMA values. We suggest that tree shelter is a promising option to restore brown oaks due to its efficacy to defend grazing and support the local community's rights to forest grazing. Our finding is expected to support Bhutan's forest policy of incorporating grazing and tree regeneration into forest management.     

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.     

Riparian Forest Restoration: Increasing Success by Reducing Plant Competition and Herbivory

The reestablishment of riparian forest is often viewed as “best management practice” for restoring stream ecosystems to a quasi‐natural state and preventing non‐point source contaminants from entering them. We experimentally assessed seedling survivorship and growth of Quercus palustris (pin oak), Q. rubra (red oak), Q. alba (white oak), Betula nigra (river birch), and Acer rubrum (red maple) in response to root‐stock type (bare root vs. containerized), herbivore protection (tree shelters), and weed control (herbicide, mowing, tree mats) over a 4‐year period at two riparian sites near the Chester River in Maryland, U.S.A. We started with tree‐stocking densities of 988/ha (400/ac) in the experimental plots and considered 50% survivorship (i.e., a density of 494/ha [200/ac] at crown closure) to be an “acceptable or minimum” target for riparian restoration. Results after four growing seasons show no significant difference in survivorship and growth between bare‐root and containerized seedlings when averaged across all species and treatments. Overall survivorship and growth was significantly higher for sheltered versus unsheltered seedlings (49% and 77.6 cm vs. 12.1% and 3.6 cm, respectively) when averaged across all species and weed control treatments. Each of the five test species exhibited significantly higher 4‐year growth with shelter protection when averaged across all other treatments, and all species but river birch had significantly higher survivorship in shelters during the period. Seedlings protected from weeds by herbicide exhibited significantly higher survivorship and growth than seedlings in all other weed‐control treatments when averaged across all species and shelter treatments. The highest 4‐year levels of survivorship/growth, when averaged across all species, was associated with seedlings protected by shelters and herbicide (88.8%/125.7cm) and by shelters and weed mats (57.5%/73.5 cm). Thus, only plots where seedlings were assisted by a combination of tree shelters and either herbicide or tree mats exhibited an “acceptable or minimum” rate of survivorship (i.e.,>50%) for riparian forest restoration in the region. Moreover, the combined growth and survivorship data suggest that crown closure over most small streams in need of restoration in the region can be achieved most rapidly (i.e., 15 years or less) by protecting seedlings with tree shelters and controlling competing vegetation with herbicides.     

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.     

Tree fern trunks facilitate seedling regeneration in a productive lowland temperate rain forest

Seedling regeneration on forest floors is often impaired by competition with established plants. In some lowland temperate rain forests, tree fern trunks provide safe sites on which tree species establish, and grow large enough to take root in the ground and persist. Here we explore the competitive and facilitative effects of two tree fern species, Cyathea smithii and Dicksonia squarrosa, on the epiphytic regeneration of tree species in nutrient-rich alluvial forests in New Zealand. The difficulties that seedlings have in establishing on vertical tree fern trunks were indicated by the following observations. First, seedling abundance was greatest on the oldest sections of tree fern trunks, near the base, suggesting that trunks gradually recruited more and more seedlings over time, but many sections of trunk were devoid of seedlings, indicating the difficulty of establishment on a vertical surface. Second, most seedlings were from small-seeded species, presumably because smaller seeds can easily lodge on tree fern trunks. Deer browsing damage was observed on 73% of epiphytic seedlings growing within 2 m of the ground, whereas few seedlings above that height were browsed. This suggests that tree ferns provide refugia from introduced deer, and may slow the decline in population size of deer-preferred species. We reasoned that tree ferns would compete with epiphytic seedlings for light, because below the tree fern canopy photosynthetically active radiation (PAR) was about 1% of above-canopy PAR. Frond removal almost tripled %PAR on the forest floor, leading to a significant increase in the height growth rate (HGR) of seedlings planted on the forest floor, but having no effects on the HGRs of epiphytic seedlings. Our study shows evidence of direct facilitative interactions by tree ferns during seedling establishment in plant communities associated with nutrient-rich soils.     

Does shelter enhance early seedling survival in dry environments? A test with eight Mediterranean species

Question: Do solid‐walled polyethylene tubes and mesh fabric tubes improve the short‐term survival of eight Mediterranean tree and shrub species often used in the restoration of arid environments? Location: We conducted two experimental plantations in degraded field sites in the province of Almería (SE Spain), under arid Mediterranean conditions. Methods: One‐year‐old seedlings of Ceratonia siliqua, Juniperus phoenicea, Olea europaea, Pinus halepensis, Pinus pinaster, Quercus coccifera, Quercus ilex and Tetraclinis articulata were planted either sheltered by one of the above shelter tubes, or by being left unsheltered. Survival was recorded the first growing season after planting, which was a very dry season. Results: Overall, seedling survival ranged from as little as 0% to 24%, and tree shelters consistently enhanced survival in Quercus species only, ranging from 16% in walled shelters to 8% in mesh shelters. Shelters failed to boost survival in the six remaining species. Conclusion: The results of this study suggest that both walled and mesh shelters were mostly ineffective at increasing seedling survival for the Mediterranean species used in this experiment; these species coincide with those used in restoration programs. The use of shelters in restoration programs conducted in arid environments should be reconsidered, while walled shelters might be advisable for Mediterranean Quercus species only. Further research is necessary to develop and assess improved types of shelters for arid environments.     

The response of forest floor vegetation and tree regeneration to deer exclusion and disturbance in a riparian deciduous forest,central Japan

The response of forest floor vegetation and regeneration of major treespecies to deer exclusion in a riparian forest were studied for 3 years withtheinteractive effects of natural disturbances. At the start of this study, deerdensity had quickly increased to a fairly high level (29–31 individualskm^–2) during the last decade and had severely reduced theamount of forest floor vegetation in the study area. Dwarf bamboos, which weredominant before, had almost diminished from the forest floor. During the studyperiod, aboveground biomass increased steadily but species diversity did notchange much in the exclosures. Outside the exclosures, the seedlings of alltreespecies were damaged greatly by deer browsing, especially the taller ones. Deerbrowsing had little effect on the emergence of tree seedlings, but deertrampling might have accelerated emergence indirectly by disturbing the soilsurface for some species. Differences in plant responses were also observed fordeer browsing and the presence of dwarf bamboo that strongly inhibits therecruitment of tree seedlings. The nine major species were classified intothreegroups according to the response of seedlings to deer browsing and disturbance.Group 1 consists of the species whose seedling survival is affected bybrowsing,but seldom by disturbances (Phellodendron amurense,Kalopanax pictus, Quercus crispulaandMalus toringo). Groups 2 and 3 consist of species adaptedto tree-fall disturbance (Betula spp.) and ripariandisturbance (Alnus hirsuta var.sibirica, Ulmus davidiana var.japonica, Populus maximowiczii andSalix sachalinensis), respectively, and seedling survivalof these two groups is principally affected by light conditions. The effect ofdeer browsing on seedling survival and growth was greater for Group 2 than forGroup 3.     

Response of young<Emphasis Type="Italic"> Tsuga heterophylla</Emphasis> to deer browsing: developing tools to assess deer impact on forest dynamics

We used dendroecology to describe and understand the consequences of deer browsing on regenerating western hemlock (Tsuga heterophylla). We compared tree shape, growth rate, height and age at four different sites in Haida Gwaii (British Columbia, Canada) that had trees representative of the range of deer impact on trees: (1) trees showing no sign of browsing, (2) escaped trees which were still browsed below the browse line and (3) stunted and heavily browsed trees. Repeated and intense browsing resulted in the small size, compact heavily ramified shape of stunted trees and in the short compact and ramified lower branches of escaped trees. These contrasted with the shape of non-browsed trees, a shape that was also found in escaped trees above the browse line. Before release, all browsed trees experienced stagnation in growth characterised by narrow rings (0.3 mm/year) and a small annual height increment (2.5 cm/year). At release, growth rate increased and stabilised: rings were wider (1.3 mm/year) and annual height increments were greater (10.5 cm/year). Non-browsed trees had a mean ring-width of 1.3 mm/year and an annual height increment of 22 cm/year. Delay in tree recruitment caused by deer varied from site to site. It had been about 15 years for the escaped trees and is estimated at 30–40 years for the stunted trees. Spatial variation in deer impact may reflect spatial variation of browsing pressure resulting from local differences in the availability of preferred forage or to differences in tree chemical defences/nutritional values.     

Effects of Tree Guards and Weed Mats on the Establishment of Native Tree Seedlings: Implications for Forest Restoration in Hong Kong, China

The effects of tree guards and weed mats on establishment and growth of native tree seedlings, Thick‐leaved oak (Cyclobalanopsis edithiae (Skan) Schott., Fagaceae), planted in an exposed hillside grassland in Hong Kong, were investigated. The natural regeneration of C. edithiae is poor due to a lack of seed dispersal agents and high seed predation, and therefore, this species is often targeted for forest restoration. The experiment lasted for 3.5 years during which the height, basal diameter, and crown diameter of individual seedlings were measured and survivorship recorded. The use of weed mats alone did not have a significant effect, but a combination of tree guards and weed mats led to a significant improvement in establishment, survivorship, and growth of the seedlings during the experimental period. Initially, the guards promoted rapid height growth of the seedlings, although lateral growth and secondary stem thickening were compromised. After the seedlings grew over the tree guards, the basal diameter and crown diameter increased at a notably faster rate. The combined effect of the tree guard and weed mat on the seedling growth pattern was found to be beneficial and contributed to the high survivorship of the seedlings. Comparing the survivorship data and the costs of various treatments, the use of tree guards in combination with the weed mats was demonstrated to be more cost‐effective than planting the seedlings without tree guards or weed mats. The potential for applying the technique in afforestation programs with native tree species for forest restoration in Hong Kong and other tropical regions is discussed.     

Influence of herbivores on a perennial plant: variation with life history stage and herbivore species

Herbivores have diverse impacts on their host plants, potentially altering survival, growth, fecundity, and other aspects of plant performance. Especially for longer-lived plant species, the effects of a single herbivore species can vary markedly throughout the life of the host plant. In addition, the effects of herbivory during any given life history stage of a host plant may also vary considerably with different types of herbivores. To investigate the effects of herbivory by black-tailed deer (Odocoileus hemionus columbianus) and snails (Helminthoglypta arrosa and Helix aspersa) on a nitrogen-fixing shrub, Lupinus chamissonis, we established three exclosure experiments in a sand dune system on the coast of northern California. These experiments documented that deer browsing significantly reduced the volume and growth rate of lupines in the seedling and juvenile life stages. Since plant volume was strongly correlated with aboveground dry biomass for lupines, such herbivore-induced reductions in volume should translate into losses of aboveground biomass. Deer browsing also significantly altered the likelihood of attack by and density of a leaf-galling cecidomyid fly (Dasineura lupinorum), suggesting that a vertebrate herbivore indirectly affected an invertebrate herbivore in this system. Although deer did not significantly affect the survival of lupine seedlings and juveniles, individuals protected from deer had consistently greater survival in the two separate experiments. Our results revealed that snails did not have a significant effect on the survival or growth of juvenile plants, despite being common on and around lupines. An exclosure experiment revealed that herbivory by deer significantly reduced the shoot lengths of mature shrubs, but led only to a minimal reduction in growth rates. In addition, we found that browsed shrubs had significantly greater inflorescence production, but also produced individual seeds with significantly reduced mass. Collectively, these data indicate that deer and snails have widely differing effects on their shared host plant; browsing by deer indirectly affects insect herbivores, and the impacts of deer change markedly with the life history stage of their host plant.     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.     

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.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Post-Planting Treatments Increase Growth of Oregon White Oak (Quercus garryana Dougl. ex Hook.) Seedlings

The extent of Oregon white oak woodland and savanna ecosystems in the U.S. Pacific Northwest has diminished significantly during the past century due to land use changes and fire suppression. Planting Oregon white oak seedlings is often necessary when restoring these plant communities. Our objective was to determine the efficacy of post‐planting treatments for establishing Oregon white oak seedlings on sites characterized by low growing season precipitation and coarse‐textured soils. We evaluated the effects of control of competing vegetation, tree shelters, fertilization, irrigation, and planting date on growth of planted seedlings. Survival was generally high (90%), but growth rate varied substantially among treatments. Plastic mulch increased soil water content and increased annual seedling height growth by an average of 56% relative to one‐time manual removal of competing vegetation. Solid‐walled tree shelters reduced browse damage and increased mean annual height growth compared to mesh shelters and no shelter by averages of 7.5 and 10.9 cm, respectively. Controlled‐release fertilizer applied at planting did not consistently increase seedling growth. Weekly irrigation (3.8 L/seedling) increased first‐year seedling growth only where mulch also was applied. Seedlings planted by late February had greater root growth by summer than those planted in early April. Soil water management was necessary for best seedling growth, and the improved height growth in solid‐walled tree shelters allowed the terminal shoot to grow more quickly above the height of animal browse. Our results indicate effective methods for establishing Oregon white oak seedlings, but these results may also be applicable to establishment of other tree species on similarly droughty sites.     

Effects of large herbivores and fire on the regeneration of Acacia erioloba woodlands in Chobe National Park, Botswana

Acacia erioloba woodlands provide important forage and shade for wildlife in northern Botswana. Mortality of mature trees caused by browsing elephants has been well documented but the lack of regeneration of new trees has received little attention. Annual growth of new shoots and changes in height were measured to determine the influence of elephants and small ungulate browsers, rainfall and fire on the growth and survival of established A. erioloba seedlings from 1995 to 1997 in the Savuti area of Chobe National Park. All above‐ground vegetation was removed from 40% of established seedlings in 1995 and 28% in 1997 by browsing elephants, and the mean height of remaining seedlings decreased from >550 mm to <300 mm. When seedlings browsed by kudu, impala and steenbok but not elephants are considered, mean seedling height increased <50 mm per year, even though mean new shoot growth remaining at the end of the dry season was 100–200 mm. Fires burned portions of the study area in 1993 and 1997, killing above‐ground vegetation, but most established A. erioloba seedlings survived, producing coppice growth from roots. While elephants and fire caused the greatest reduction in established seedling height and number, small browsers suppressed growth, keeping seedlings vulnerable to fire and delaying growth to reproductive maturity.     

How do deer affect tree seedlings on a dwarf bamboo-dominated forest floor?

A field experiment was conducted in Mt. Ôdaigahara, west-central Japan, to reveal the effects of two herbivores, Sika deer (Cervus nippon) and mice (Apodemus spp.), on the seedlings of five cohorts of three tree species, Abies homolepis, Fraxinus lanuginosa f. serrata and Fagus crenata. The forest floor of the study site was covered with dwarf bamboo, Sasa nipponica, which would also affect the seedlings. Eight combinations of three treatments were set: exclusion of deer, exclusion of mice and removal of dwarf bamboo. Deer were expected to affect the seedlings not only negatively by foraging but also positively by browsing dwarf bamboo that overshadows seedlings. The survival of these cohorts was analyzed by survival analysis and the differences in their survival between the treatments were investigated. The results of the experiment showed that: (1) exclusion of deer increased the aboveground biomass of dwarf bamboo and made it more difficult for seedlings to survive under the regenerated dwarf bamboo stand, and (2) deer negatively affected the seedlings, but they had positive, indirect effects in some cohorts by decreasing the aboveground biomass of dwarf bamboo. However, such effects were not detected in some cohorts which were affected excessively by deer or dwarf bamboo. No clear results on the effect of mice on seedlings were obtained. We conclude that a positive, indirect effect of deer on seedling survivorship would be observable when the magnitude of a positive, indirect effect caused by decreasing dwarf bamboo is larger than that of a negative, direct effect of deer.     

Ungulate browsing on introduced pines differs between plant communities: Implications for invasion process and management

The effects of herbivory on plant invasions are broadly discussed, and many studies have led to widely debated theories. In particular, the effects of herbivores on pine invasion found in different studies vary; in some cases, they controlled their expansion, and in others, they promoted it. On the other hand, vulnerability to invasion by pines differs between community types. Sites with dunes and bare ground are the most heavily invaded, followed by grasslands, while shrublands and forests are least invaded. Because current evidence is mostly observational, some of the varying responses of pine invasions to herbivory should be examined further through replicated experiments. Here, we address experimentally the extent to which preference for the non‐native invasive Pinus contorta by domestic sheep (Ovis aries) depends on the vegetation type. We installed experimental enclosures within two adjacent communities, grassland and shrubland, and in each one, we planted seedlings of P. contorta Douglas and established a sheep density typically recommended for the study area. The number of browsed seedlings, the number and type of branches browsed per seedling, the reduction in height and probability of survival immediately after browsing period were recorded. The number of browsed seedlings and damage to the terminal bud were higher in grassland than in shrubland, while the number of browsed branches per seedling was higher in shrubland than grassland. The reductions in height and probability of survival immediately after browsing were similar in both communities. These results show that moderate levels of sheep herbivory could reduce 20% seedling survival in both communities; nevertheless, the damage patterns differ between them. The sheep browsed more substantial number of seedlings in grasslands than in shrublands. However, if sheep find the seedlings, they damage it more in shrublands. These results suggest that experimental studies comparing communities are important for pine invasion management.     

Patterns of woody plant abundance,recruitment, mortality,and growth in a 65 year chronosequence of old-fields

We surveyed vegetation along forest margins in a 65-year chronosequence of old-fields at the Cedar Creek Natural History Area in east-central Minnesota, USA, to identify successional patterns of woody plants and to determine if these were correlated with soil nitrogen. We predicted that shrub and tree abundance, size, and distance of occurrence from the forest edge would be correlated with field age or soil nitrogen. Instead we did not find successional trends in the abundance or composition of woody species. Even in the oldest field the abundance of trees and shrubs was low and concentrated in areas close to the forest. Though trees were larger and present further from the forest edges in older fields, average tree height was less than 126 cm in all fields.Since we did not find successional trends we looked at various local factors (local seed sources, deer browsing, and forest edge aspect) and their relation to recruitment, mortality, or growth to explain variation among fields in abundance of trees or shrubs. The three most common tree species (Quercus rubra, Q. macrocarpa,and Populus tremuloides) all had a higher relative abundance of seedlings, and two (Q. rubra and Q. macrocarpa) had a higher relative abundance of large trees adjacent to forests with a high abundance of conspecific adults. Most trees taller than 20 cm were browsed by deer and were shorter in 1995 than they were in 1993. Mortality was higher for trees less than 30 cm indicating that mortality was size-dependent. Forest edge aspect did not significantly influence the abundance or demography of any species. Our results suggest that the patterns of seedling recruitment were largely determined by the proximity of seed sources and that these patterns may persist so that tree communities in old-fields resemble adjacent forests. Deer may be a significant factor in the suppression of tree populations in old-fields through repeated browsing which reduces tree growth and elevates tree mortality by prolonging the period of time trees remain susceptible to size-dependent mortality.     

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.     

Effects of deer,mice and dwarf bamboo on the emergence,survival and growth of <Emphasis Type="Italic">Abies homolepis</Emphasis> (Piceaceae) seedlings

In the subalpine mixed forest of Mt. Ôdaigahara, mid-western Japan, the understory is dominated by dwarf bamboo (Sasa nipponica), which is the major forage of overly populous sika deer (Cervus nippon). In the present study, we monitored the survival and growth of Abies homolepis seedlings over 5years to determine how they responded to the experimental removal of dwarf bamboo and to the exclusion of sika deer and mice (Apodemus argenteus and A.speciosus). Deer and dwarf bamboo reduced the survival of seedlings but had different effects on growth. The stems of seedlings were shorter in the presence of deer, indicating that taller seedlings were apt to be browsed by deer, whereas the diameters of seedlings were smaller in the presence of dwarf bamboo, mainly owing to its shading effect. The presence of mice decreased the number of seedlings germinating in a particular site, but had no effect on seedling survival after germination. There was no significant indirect effect whereby the survival of seedlings was predicted to be facilitated by the decreased biomass of bamboo because of grazing by deer. We supposed that this might be because the direct negative effect of deer was so large as to conceal the positive indirect effect.