Leader Browsing by Red and Roe Deer on Young Sitka Spruce Trees in Western Scotland. II. Effects on Growth and Tree Form

Some 2000 trees at 14 main sites in Glenbranter Forest, Argyll,were monitored during 1978–1990 for damage. Tree state(whether leaderless or with single or multiple leaders or trunks)and tree size were regularly recorded. Browsing of leaders depressed height increment by 5–10cm in the year of ocurrence compared with undamaged trees; therapid development of new leaders minimized the check in growth.Trees initially leaderless had equal height increment over ayear to trees initially with leaders if no damage occurred inthe year. At a subsidiary site where some trees were planted in smallexclosures, the trees protected from deer reached a safe heightafter five seasons of growth, unprotected trees reaching equalheight 1 year later. The girths attained when trees were 10–15 years old wereon average only slightly depressed by earlier browsing damageto leaders, reductions at most equalling one year's increment. Many trees became multi-trunked, there being a significant relationshipbetween the percentage incidence of multi-trunking and the numberof occurrences of leader damage. Unbrowsed trees mostly becamesingle-trunked. We judge that the present high incidence ofmulti-trunking (45 per cent in our 9–15 year-old stands)results largely from deer browsing.     

Patterns of Stem Damage by Moose (Alces alces) in Young Pinus sylvestris Stands in Sweden

The incidence of recent moose browsing of the main stem on young Scots pine (Pinus sylvestris L.) trees, and correlations with tree or stand characteristics, were examined using data from a large-scale survey in Sweden. On average, 10% of all Scots pine main stems showed one or more types of recent damage, with browsing of the apical leader accounting for about 75% of all damage recorded. Stripping of bark off the main stem occurred most frequently in the southern area, where site productivity is highest. Apical leader browsing was negatively correlated with height of the main stems, length of the apical leader and incidence of pre-commercial thinning, whereas bark stripping and stem breaking were positively correlated with the same variables. All types of recent stem damage correlated positively with the extent of previous stem damage and also with the extent of recent browsing of lateral shoots.     

Height growth of young larch (Larix kaempferi) in relation to the frequency of deer browsing damage in Hokkaido, Japan

The effect of browsing frequency by Sika deer (Cervus nippon yezoensis) on the height growth of Japanese larch (Larix kaempferi) in Hokkaido, Japan, was analyzed. Tree heights at the time of the census and at the start of the current growing season were determined for 100 trees selected randomly at each site. The tallest shoot on each tree was identified and the number of browsing occurrences on the current-year shoot was determined. Young larches browsed by deer repeatedly develop new shoots during a growing season. Larch is a browsing-tolerant species, and the average height growth of the most heavily browsed trees was >15 cm. Although resprouting partly compensated for decreased height growth caused by browsing, the annual increase in height decreased with increased browsing frequency. Thus, browsing frequency during a growing season best reflects the intensity of browsing damage to larch trees.     

Timing, light availability and vigour determine the response of Abies alba saplings to leader shoot browsing

Herbivore browsing on tree saplings is a common phenomenon that can cause damage particularly on preferred species. In this study, the combined effects of light availability and timing of browsing on the response of 9-year-old Abies alba saplings were tested experimentally. Leader shoot clipping was applied before budburst, shortly after budburst or in autumn on saplings grown in full light or under artificial shade. Timing of clipping, light availability and tree vigour (expressed as height and tree ring width before clipping) had an effect on the height after clipping. After clipping in autumn or before budburst, fast-growing fir saplings bent up twigs to form new leader shoots and overcompensated height loss; saplings characterised by intermediate growth rates formed new shoots out of regular visible lateral buds; and slow-growing saplings had no new shoot in the first year after clipping, such that the clipping-induced height difference even increased over time. Saplings clipped shortly after budburst elongated the remaining part of the shoot in the first year and developed shoots out of the most distal lateral buds in the second growing season, leading to complete height compensation. Multi-trunking was typical for all clipped trees. We conclude that the microscale conditions under which a tree is growing (i.e. which affect tree vigour) are highly important for determining whether the height reduction imposed by browsing is offset by overcompensation or increases over time relative to unclipped trees. This response can partly be influenced by forest management via enhancing tree vigour via the light regime.     

Effects of deer abundance on broad-leaf tree seedling establishment in the understory of <Emphasis Type="Italic">Abies sachalinensis</Emphasis> plantations

Browsing by sika deer (Cervus nippon) has significant negative effects on regeneration in many forests in Japan; however, the effects of browsing on regeneration processes have not been determined quantitatively. Our study was conducted in Abies sachalinensis plantations in seven tracts with differences in deer abundance on Hokkaido, northern Japan, to identify indicators for the effects of deer on the regeneration of broad-leaf species from observing seedlings. Five 5 × 20-m plots were located within each tract, and the densities of seedlings 30–200 cm tall and percentages of browsed seedlings were determined. We used sightings per unit effort (SPUE) by hunters and spotlight survey counts (SLCs) as indices of deer abundance for each tract. Seedling density was negatively correlated with deer abundance and coverage of dwarf bamboo, and basal areas of overstory trees also affected number of seedlings. Percentage of browsed seedlings was positively correlated with deer abundance, and it was affected by deer preferences among seedling species. However, the percentage of browsed seedlings was more clearly related to deer abundance than seedling density. There were few seedlings of tree species ≥100 cm tall in tracts with the highest deer abundance. Based on these results, deer abundances of SPUE >6 sightings per hunter-day or SLC >15 animals per 10 km are likely to prevent regeneration of broad-leaf species. The percentage of browsed seedlings and density and browsing damage on tree seedlings ≥100 cm tall are useful indicators of the effects of deer.     

Browsing damage on broadleaved trees in semi-natural temperate forest in Sweden,with a focus on oak regeneration

Absence of, or poor, oak (Quercus spp.) regeneration is a problem in uneven-aged, mixed closed-canopy broadleaved forests. Browsing by ungulates on small trees may contribute to poor oak regeneration in such forests. This possibility was investigated in 25 Swedish stands, and browsing damage was analysed in relation to landscape and stand factors. The proportion of browsed small (<20 cm tall) oak seedlings and other seedlings was low, and apparently a minor mortality factor. For saplings (20–130 cm tall), accumulated browsing damage was generally higher on oak than on five major competing tree species: Fraxinus excelsior, Corylus avellana, Tilia cordata, Acer platanoides and Sorbus aucuparia. Leaf removal was rare in late summer, except for rowan. The amount of cover (shelter) for ungulates near plots was positively correlated with oak browsing intensity; within plots, a high density of ash saplings may reduce browsing on oak saplings. In these forests, browsing probably retards growth of oak saplings relative to competing trees. Oak may persist as a minor stand component, but monitoring is needed to study future changes.     

Browsing and damage inflicted by moose in young Scots pine stands subjected to high-stump precommercial thinning

In Fennoscandia, young stands of Scots pine (Pinus sylvestris L.) are intensively used by moose (Alces alces L.) during winter. We studied whether forage amounts on high-cut pines in high-stump commercial thinning influenced browsing intensity and damage incidence on retained (i.e. uncut) pine stems. High-cut pines were browsed, but to a lesser extent than retained pines. At a scale corresponding to individual feeding sites (≈40 m²), browsing intensity on retained pines was not influenced by the amount of forage on high-cut pines but was positively related to moose pellet group counts. The incidence of lower-height damage (stem breakage and bark stripping) was positively related to the amount of forage on high-cut pines, whereas higher damage (leader shoot browsing) was not. Overall browsing damage incidence on retained pines was positively related to the density of deciduous trees and negatively related to the amount of forage on retained pines. Our results suggest that although high-stump thinning supplies additional food resources for moose, larger amounts of forage on high-cut pines may increase the risk for bark stripping and stem breakage on retained trees. Further research is needed at larger spatial scales to assess the feasibility of high-stump thinning as a damage mitigation measure.     

Growth of Young Sitka Spruce (Picea sitchensis (Bong.) Carr.) and the Effect of Simulated Browsing, Staking and Treeshelters

Sitka spruce (Picea sitchensis (Bong.) Carr.) 1 + 1 transplantswere grown at 1 x 1 m for 3 years with 100 per cent chemicalweed control on a very fertile site. Treatments were control,firm staking, staked with 15 cm diameter netting with simulatedbrowsing, staked with simulated browsing to 15 cm diameter crown,1.2 m treeshelter, 1.2 treeshelter with holes and no browsing.No leaders were browsed. Trees were assessed for annual heightgrowth, stem diameter at 0.5 m and 1.0 m, total above and belowground biomass, root:shoot ratio, root plate diameter, rootdepth and root volume. No 3-year heights were significantlydifferent but third-year increments were; the greatest finalheight was in the control at 200.7 cm. All other tree parametershad significant treatment differences demonstrating the needto select carefully the criteria of success in experiments.Treatments which restricted the horizontal development of thecanopy either physically or by simulated browsing seriouslyreduced stem diameter and root development. The controls hada root:shoot ratio of 0.418; treeshelter treatments had thesmallest root:shoot ratio of 0.238; staked trees were 0.379while staked browsed trees had the highest value at 0.447. Acceptanceof browsing or use of shelters or guards to protect trees frombrowsing may seriously reduce growth.     

Simulated effects of browsing on the production and economic values of Scots pine (Pinus sylvestris) stands

Effects of browsing damage on Scots pine (Pinus sylvestris) production and economic value were simulated on sites ranging in fertility from relatively poor to relatively high. Effects of browsing damage on growth and survival after trees had exceeded browsing-height were estimated using data from a browsing study. Stem lists were produced when stems had reached a height of about 4?m. Individual tree data were then imported into the Heureka simulation system to generate predictions about stand production and economic value. Six browsing levels were simulated: 0, 2, 5, 10, 15 and 27% browsing damage annually. The 2% and 5% browsing levels had only minor effects on production, whilst higher browsing levels significantly reduced production. However, the effect on economic value was more pronounced and even the 2% browsing level had significant effects on estimated land expectation values. Results from this simulation may support discussions about acceptable browsing levels in young Scots pine stands.     

Establishment patterns of woody species in low intensity-grazed pastures after the cessation of intensive agricultural use

Low intensity grazing with large herbivores is increasingly used to steer the development of mosaic landscapes (including wood-pastures) and woodland expansion on former agricultural land. We studied the establishment and early growth pattern of woody species in grasslands (formerly used for high intensity summer cattle grazing) in which a low intensity-grazing regime was set up between 4 and 11 years ago. We tested whether the developing vegetation patches, consisting of tall herbs and scrub, influenced this pattern. Further, we investigated if increased susceptibility to trampling disturbance by large herbivores on wet soils in floodplains influenced sapling establishment.Formerly intensively used grasslands were studied at six sites, four in floodplains and two in interfluvial zones on nutrient rich, heavy soils. In total, we sampled 141 plots with grassland, tall herb or scrub vegetation and recorded individuals of established tree and shrub species, their height, diameter, browsing damage and distance to potential seed sources. In the floodplains, we recorded soil disturbance by large herbivores (hoofprints) in wet and moist soils.Sapling frequencies, with Fraxinus excelsior as the most common species (>50% of all individuals), were two to three times higher in tall herb and scrub patches compared to grassland. Only the spiny Crataegus monogyna established in higher frequencies in grassland plots. In floodplains, we found two to three times more established woody saplings in wet tall herb plots, compared to moist and wet grassland plots and moist tall herb plots.Browsing reduced sapling height in grassland and tall herb patches. Only in scrub patches did average height (±2.5 m) reach above the browse line. We found 30–70% less browsing damage to saplings in scrub compared to damage in grassland and tall herb plots. In particular, damage to the apical shoot was strongly reduced.Our findings show that tree establishment occurs in spatial association with non-grassland patches that offer protection against browsing. Fast growing spiny shrubs like Rubus sp. offer protection for palatable saplings, allowing establishment and growing out beyond the browse line. Alternatively, increased establishment occurs in wet tall herb patches, where soil disturbance by moderate trampling favours germination and, subsequently, saplings are protected at least temporarily against browsing by unpalatable tall herbs.     

Height growth recovery and crown development in top‐damaged Pinus sylvestris trees

Leader shoot damage was imposed on young Scots pines (3 m high) in four ways: by caging the uppermost whorl including the leader shoot with 10 pine shoot beetles; by caging the two uppermost whorls with 20 beetles; by topping the tree below the 1984 whorl; by topping the trees as above and removing all current shoots in whorl 1983. Six years later, top‐damaged trees were still significantly shorter than undamaged controls, although the annual height growth rate had recovered to the pre‐damage level within 3 years. Recovery occurred in two ways: lateral branches from the nearest intact whorl took dominance over the damaged leader or secondary branches developed from interfascicular buds on the damaged leader shoot. Approximately one and two years of height growth were lost in the former and latter case, respectively. Only 5 of the 34 top‐damaged trees escaped technical defects (crooks and/or vertical branches). Stem diameter and crown development were slightly affected.     

Deer Browsing and Impact on Forest Development

Abstract

Browsing by large herbivores can cause serious costs in forest production. The most important costs related to browsing damage are insufficient stand regeneration, volume losses and timber quality impairment. In Sweden, moose (Alces alces)and roe deer (Capreolus capreolus) are the most abundant deer species and therefore cause the majority of browsing damage. Balancing costs and benefits related to deer density is difficult since few data about the final losses in timber volume or quality impairment is available. Simulations of browsing and the impact on stand growth suggest that a minor or moderate browsing intensity has little effect on final harvest volume. The reason is that trees seem to recover some of the lost growth as they grow out of reach of the animals. Balancing between different uses of the forestry resource is complicated by our limited capability to predict the impact of different management regimes.     

Conifer leader browsing by roe deer in English upland forests: Effects of deer density and understorey vegetation

At moderate to high densities ungulates can impact negatively on forest crops and these may be managed by lethal control. In production forestry an understanding of the relationship between ungulate density, habitat-related factors and the incidence of tree damage may promote more efficient ungulate damage management than by lethal control alone. In plantation forests in the north east of England, the incidence of conifer leader browsing by roe deer (Capreolus capreolus) was positively associated with an index of deer density (at the site scale) and planted crop tree density (at the feeding patch scale) and was negatively associated with the relative abundance of palatable understorey vegetation (at the site scale). An interaction between the deer density index and palatable understorey browse abundance was positively associated with leader browsing. Coefficients from the optimal model were used to construct a browse probability surface across a matrix of values of deer dung density and palatable understorey vegetation abundance. This illustrated a very different relationship between dung density and conifer leader browsing probability across the various levels of palatable understorey vegetation abundance. From this we hypothesise that in areas of limited palatable understorey vegetation abundance, higher levels of culling may be required to achieve reductions in the incidence of conifer leader browsing than would be necessary in areas with moderate to high palatable understorey vegetation abundance. Improvements to understorey vegetation in concert with roe deer population control may enhance the efficiency of deer damage management in upland areas with nutrient-poor soils.     

Bark Stripping Damage by Red Deer in a Sitka Spruce Forest in Western Scotland: II. Wound Size and Position

Wounds to bark were measured within a few months of being inflictedby red deer in Glenbranter Forest, Argyll. About 8000 treeson 240 permanent plots were monitored; all the wounds occurringover a period of seven years were examined. Wound length varied little between forest stages, averaging12 cm. Wound breadth was greater on older trees, averaging 5cm in Sitka spruce and 6 cm in Norway spruce for the pole andhigh-canopy^* forest stages. A few wounds were large, maximumlengths being 95 and 80 cm in Sitka spruce and Norway sprucerespectively, but less than 20 per cent of the Sitka sprucewounds exceeded 100 cm² in surface area. Seasonal variationin size was modest; wounds were longest in spring (April–June). Often a single tree received two or more wounds during the 3-or6-month-long observation periods. For Sitka spruce there were89 instances of multiple wounding and 142 of single wounding,but these single wounds contributed only 37 per cent of thetotal of 383 wounds observed. The area of trunk wounded wasmore dependent on the size of the largest wound than on woundnumber, multiple wounds often being small. Trunks received most damage between 50 and 100 cm above ground,the lower and upper limits of wounding being 2 and 150 cm. Thepeaking in the incidence of damage with regard to height wasmost marked in pole-stage forest, whereas wounds were spreadmore evenly up the trunks on younger trees. Wound height waslittle affected by season, mean basal heights being 68, 70,74 and 67 cm in winter, spring, summer and autumn. Woundingon the trunk was not correlated with aspect.     

Effects of shelter tubes on hardwood tree establishment in western Oregon silvopastures

Honey locust (Gleditsia triacanthos), black locust (Robinia pseudoacacia), and honey mesquite (Prosopis glandulosa) are warm season forage trees with potential to efficiently share site resources with cool season pasture plants in Pacific Northwest silvopastures. Establishment of hardwood trees can be difficult, however, because of feeding damage from wildlife and livestock. This study compared establishment and growth of trees planted in 88 cm tall solid plastic shelter tubes to 88 cm tall plastic mesh tubes, used to protect trees from animal damage. Three replications were established in May 1995 for each of the three tree species on a hill pasture near Corvallis, Oregon. Initial tree survival during the first summer and winter following planting was higher in shelter tubes than in mesh tubes. At the end of the third growing season, 58% of black locust and 94% of honey locust trees in shelter tubes were still alive compared to only 14% of black locust and 47% of honey locust in mesh tubes. Few honey mesquite trees survived regardless of tube type used. Average three-year total height growth for black locust was increased by 650% and basal diameter growth by 380% within shelter tubes, while honey locust height growth was increased by 300% and diameter growth was increased by 150% compared to trees in mesh tubes. However, shelter tube trees tended to be taller relative to their diameter and had difficulty standing upright if tubes were removed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Influences of seedling size,container type and mammal browsing on the establishment of <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> in plantation forestry

Rapid early growth of tree seedlings is critical to the success of plantation establishment. We investigated the effects of seedling size (small and large) and container types (small [Lannen 121], medium [Lannen 81] and large [Forestry Tube]) in the nursery and the effects of mammal browsing after planting on growth of Eucalyptus globulus in Tasmania’s Southern Forests. After planting, seedlings were either exposed to browsing or protected from browsing by wire-mesh cages until age 6 months. Low browsing pressure resulted in around 20% and 5–10% of foliage being browsed in the large and small size categories, respectively, between 1 and 3 months after planting the uncaged treatment. 6 months after planting, height growth increment was lower, and 4 years after planting, mortality was higher in uncaged large than caged large seedling treatments. Six and twelve months after planting, seedlings raised in Forestry Tube containers had significantly greater height increment and root collar diameter relative to other treatments. By 4 years after planting, trees of the small seedling treatment had significantly greater diameter than those of the large seedling treatment, but there was no effect of container type treatment. Four years after planting there was no effect on diameter growth by browsing of <30% of foliage up to 3 months after planting, although there was greater incidence of double leaders in trees that had been browsed as seedlings. Small seedlings produced more growth 4 years after planting than large seedlings of E. globulus.     

Growth change of young Picea sitchensis in response to deer browsing

Taking advantage of the introduction of the black-tailed deer to the Queen Charlotte Islands (British Columbia, Canada), we used dendrochronological analyses to understand the consequences of deer browsing on Sitka spruce growth. We compared shape, radial growth, height growth and age of young spruce in three sites. We identified two types of trees growing side by side: (1) stunted and heavily browsed spruce, smaller than the browsing limit and (2) escaped spruce that were taller than the browsing limit but still browsed in their lower part. The compact and heavily ramified shape in stunted spruce was the result of repeated and intense browsing. In escaped spruce this was also the case below the browsing limit (1.16 m±0.07 m), in sharp contrast with the normal shape that escaped spruce resumed above the browsing limit. We show that the release of browsing pressure, once the tree reaches the browsing limit, is characterised by an abrupt increase in radial growth. Before release, trees show a growth stagnation characterized by narrow rings (0.5 mm per year) and small annual height increments (<5 cm per year). After release, trees show a growth stabilisation characterised by wider rings (3 mm per year) and larger annual height increments (20 cm per year). We use this pattern to estimate frequency and age at release and their possible variation over time. Age differences between stunted and escaped spruce are highly significant and indicate that, despite of browsing, most if not all trees will ultimately reach the browsing limit and escape. Heavy deer pressure (30 deer per km²) delays spruce sapling recruitment by about 8 years. This delay varies in relation to site quality and seems to have increased over time, suggesting an increase in browsing pressure.     

Characteristics of lowland broadleaved woodland being restocked by natural regeneration

A survey of 78 sites in southern England with approved managementplans for restocking by natural regeneration was made duringthe summers of 1993 and 1994. The following features were recorded:species, stocking, canopy cover and seed-bearing potential oftrees present in the overstorey; species and canopy cover ofthe understorey; ground cover; species, browsing damage, numberand heights of tree seedlings. In general, sites were poorlystocked with overstorey trees having an average of 135 stemsha^–1 and a mean canopy cover of 36 per cent. Similarly,the understoreys were poorly developed with an average coverof only 23 per cent. Twenty-nine species of tree were foundin the overstorey, Quercus spp. and Fraxinus excelsior werethe most common. Many of the stems present had poor seed-bearingpotential. Cover of the ground flora often exceeded 75 per cent.Seedlings were present on most sites, with F. excelsior andBetula pendula being most abundant with mean seedling densitiesexceeding 10000 ha^–1. Most seedlings were >20 cm tall,few exceeded 120 cm. About 30 per cent of seedlings were browsed.The results are related to current guidance and the future prospectsfor use of natural regeneration.     

Direct impacts of sheep upon Douglas-fir trees in two agrosilvopastoral systems

Livestock may provide important service and production functions in agroforestry systems. However, use of livestock in conifer/improved pasture agrosilvopastoral systems is currently limited by concerns about potential damage to trees by livestock. Effects of sheep grazing on Douglas-fir (Pseudotsuga menziesii) trees in two patterns of sheep/pasture/conifer agroforest (cluster and grid plantations) were studied from 4 years after planting (1983) until the first precommercial thinning at age 10 years (1988). Trees averaged over 1 m in height when grazing began in summer 1983. Some browsing of tree lateral branches by sheep occurred regardless of grazing season in 1983–1985. However, the 2 to 10% of current year's lateral branch growth removed by grazing sheep was too low to impact tree growth. Sheep removed the terminal leaders from only 3 to 9% of trees each year during 1983–1985. Most browsing of terminals occurred in the summer when other forages had become mature and were relatively unpalatable to sheep. Less than 13% of agroforest trees were debarked by sheep each year during 1983–1987. By the end of grazing in 1987, less than 8% of agroforest trees had sustained a level of debarking likely to impact future growth (>50% of tree circumference debarked). Grazing had no discernible effect upon tree diameter or height in any year (P > 0.05). Total tree mortality attributable to sheep grazing during 1983–1987 was only 0.9%, including three trees girdled by sheep and two debarked trees which were subsequently attacked by insects. Overall, grazing had no detrimental impact on timber stand growth or mortality.Submitted as Oregon Agricultural Experiment Station Technical Paper No. 9628.     

Gall density of Retinia resinella (Lepidoptera,tortricidae) in relation to pine stand development

Patterns of host plant utilization of the pine resin gall moth Retinia resinella in a young Scots pine stand were studied along a belt transect in southeastern Finland. The gall distribution in 1984–1989 was retrospectively analysed. The moth preferred leaders to lateral shoots. In 1984, 60% of the galls occurred on the leaders, while the respective proportion was 40% in 1986 and 38% in 1988. The mean height of pines attacked by the moth changed from 74 to 127 cm. In 1984, moth galls on leaders occurred on higher and better‐growing trees than the average, while the opposite was true in 1988. There was an indication of clumping of R. resinella, with clumps spaced about 20 m apart which may be‐due to microhabitat distribution, the behaviour of ovipositing females or spatial dynamics of predators. The galls occurred more frequently on trees with a previous year's gall, but this tendency was not observed in the leader galls. The results support the view that the pattern of host plant utilization by a herbivore may profoundly depend on the successional stage of the habitat.