Ungulate herbivory of regenerating conifers in relation to foliar nutrition and terpenoid production

Ungulate browsing greatly influences regeneration dynamics of some forest ecosystems, yet the relationship between browse susceptibility and foliar chemistry of forest tree seedlings is not well understood. We applied field fertilization (15N-9P-10K controlled-release fertilizer at 0, 20, 40, and 60 g per seedling) and investigated how subsequent changes in terpenoid production and foliar nutrition influence ungulate browse preference for Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco), western hemlock (Tsuga heterophylla Raf. Sarg.), and western redcedar (Thujaplicata Donn ex D. Don) seedlings across four sites in northwestern Oregon, USA. Fertilization increased foliar N concentration of all three species, but above-ground growth of only Douglas-fir and western hemlock. Foliar monoterpene concentrations for western hemlock and western redcedar also increased at higher fertilization rates, while Douglas-fir monoterpene production was not affected by fertilization. Regardless of monoterpene levels, ungulate browse preference was greater for fertilized western hemlock seedlings. The opposite response, however, was observed for western redcedar at two of the four study sites where the likelihood of browse was greater for non-fertilized than fertilized seedlings. Differences in browse preference among species may depend on the type and amount of individual monoterpenes manufactured in response to fertilization. Western redcedar produce α- and β-thujone, oxygenated monoterpenes known to promote gastroenteritis and possibly inhibit microbial rumen activity, which were absent from Douglas-fir or western hemlock. Higher concentrations of α- and β-thujone associated with increasing fertilizer rate provide a plausible explanation as to why ungulates preferred non-fertilized western redcedar. Our results illustrate species-specific adaptation in browse avoidance and selective ungulate browsing behavior of individual trees as linked to foliar chemistry.     

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.     

Interactions among forestry regeneration treatments,plant vigour and browsing damage by deer.

In a large field experiment we studied the influence of regenerationmethods on the extent to which roe deer (Capreoluscapreolus L.) browse on spruce (Picea abies L.Karst) seedlings. Our objective was to evaluate if treatments that are good atstimulating regeneration may increase browsing damage by deer. Data werecollected between 1993 and 1995, and in the winters of 1993–94 and1994–95 the frequency of browsed seedlings was 13.2% and 13.4%,respectively. The frequency of browsing damage varied considerably betweensitesand clearcuts. Browsing damage was more frequent when the regeneration methodsincluded insecticide treatments. Among insecticide-free treatments, browsingwasheavier on scarified plots than on herbicide-treated or control plots (in whichseedlings were planted on untreated ground and given no post-plantingtreatment). Containerised seedlings were browsed more than bare-rooted ones.Neither the age of clearcut when planting, nor removal of slash, had any effecton browsing. It was demonstrated that regeneration methods that increased plantvigour (as assessed by leader growth and needle colour index) led to morebrowsing damage. Thus, our results support the plant vigour hypothesis, whichstates that many herbivores prefer to feed on vigorous plants because they aremore nutritious. However, the difference in browsing damage betweencontainerised and bare-rooted seedlings could not be explained solely by theplant vigour hypothesis.     

Deer prevent western redcedar (Thuya plicata) regeneration in old-growth forests of Haida Gwaii: Is there a potential for recovery?

The current increase in deer populations in many forests has fostered a growing concern about their impact on forest ecology. Sitka black-tailed deer (Odocoileus hemionus sitchensis) were introduced to Haida Gwaii (British Columbia, Canada) in the late 19th century, and they have dramatically affected the regeneration of woody species in both old- and second-growth forests since then. The lack of recruitment in western redcedar (Thuja plicata) in old-growth forests has been attributed to deer. The objectives of this study were to (1) experimentally confirm that deer browsing causes a lack of western redcedar recruitment and (2) assess the potential for and speed of recovery after a prolonged exclusion of deer. We installed a set of 20 enclosures and monitored them over a period of 8 years from 1997 to 2005. We compared temporal changes in redcedar cover and in the survival and growth of marked seedlings in plots that were or were not accessible to deer. Redcedar cover in the vegetation layer accessible to deer was generally low (from 3 to 5%) but higher inside the enclosures (an average difference of 2.3%). Protected seedlings survived better, were higher, presented more leafed shoots, and had less stems than unprotected individuals, features that suggest that deer were key to the lack of regeneration. However, growth was very slow (protected marked seedlings grew 2.5 cm on average in 8 years) and, under the current conditions, the time required for a protected seedling to escape deer would probably take over two decades. This very slow growth rate under closed canopy conditions probably reflects a gap-phase regeneration strategy and/or sensitivity to competition with other woody species. The combination of a very slow growth with a high palatability and a lack of physical defences, in contrast to the other dominant conifers in this ecosystem, probably explain why redcedar regeneration can be eliminated from old-growth forest by abundant deer populations.     

Belowground competition from overstory trees influences Douglas-fir sapling morphology in thinned stands

We evaluated effects of belowground competition on morphology of naturally established coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) saplings in 60- to 80-year-old thinned Douglas-fir stands in southwestern Washington. We separately quantified belowground competition from overstory and understory sources using trenching and understory removal. In this light-limited environment of 26 ± 16% (std. dev.) full sunlight, 2-year exclusion of tree root competition by trenching increased sapling stem biomass by 18%, total aboveground biomass by 21%, number of interwhorl buds by 68%, total foliar biomass by 33%, and foliar biomass on branch components over 4 years old by 143%. Belowground competition did not influence shoot:root ratio or foliar efficiency (i.e., stem growth per unit foliage biomass). Sapling needle size, specific leaf area, and internodal distance also were not affected by belowground competition; these variables were apparently a function of the low-light environment. The principal source of belowground competition was roots of overstory trees; effects of belowground competition from understory vegetation were minor. Thus, under a partial overstory, morphology of Douglas-fir regeneration was influenced by both belowground and aboveground competition from overstory trees. In this environment, understory vegetation control would not likely influence belowground competition to an extent that would affect sapling morphology.     

Growth reduction in harvest-age,coniferous forests with residual trees in the western central Cascade Range of Oregon

To evaluate the relationship of overstory residual trees to the growth of unmanaged young-to-mature understory Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) and western hemlock (Tsuga heterophylla (Raf.) Sarg.), the basal area and volume of 14 paired plots with and without residual trees were examined in the Willamette National Forest, Oregon. Residual trees were large survivors of the fires that initiated the understory between 55 and 121 yr ago. Understory stands were naturally regenerated and not managed in any way. High residual tree and understory densities were negatively associated with understory volume. The relation of density of residual trees to total understory and Douglas-fir basal areas and volumes was best described by a negative logarithmic function. The rate of decrease in total understory and Douglas-fir basal areas and volumes per individual residual tree became smaller with increasing residual-tree density. Predicted total understory volume reduction was 23% with five residual trees/ha and 47% with 50 residual trees/ha, averaging 4.6% and 0.9% per residual tree, respectively. After including the estimated volume growth of residual trees since initiation of the understory, stand volume was still 19% lower with five residual trees/ha and 41% lower with 50 residual trees/ha than in stands with no residual trees, averaging a reduction of 38% and 0.8% per residual tree, respectively. In mixed stands of Douglas fir and western hemlock, predicted Douglas-fir basal area and volume declined more rapidly than did total understory basal area and volume when residual-tree densities exceeded about 15 trees/ha. This difference was probably due to the relative shade-intolerance of Douglas fir. Predicted Douglas-fir volume reduction was 13% with five residual trees/ha and 75% with 50 residual trees/ha, averaging 2.6% and 1.5% per residual tree, respectively. The southern aspects had more than 150% the total understory basal area and volume and more than 200% the Douglas-fir volume and basal area of the northern aspects. Lower density and basal area of understory trees, particularly of dominant and codominant Douglas fir, were associated with increasing residual-tree densities. Given the same diameter at breast height (DBH), heights of Douglas fir were not related to residual trees. Regardless of understory age, understory volume was greatest in stands with the lowest understory densities. These results suggest that timber production in unthinned green-tree retention units may be reduced and may depend on the density of leave-trees. Thinning of understory trees is recommended to reduce growth loss from intraspecific competition.     

Can the impact of deer browsing on tree regeneration be mitigated by shelterwood cutting and strip clearcutting?

Silvicultural treatments creating large canopy openings failed to restore regeneration of balsam fir (Abies balsamea (L.) Mill.) due to browsing pressure from white-tailed deer (Odocoileus virginianus Zimmermann). Consequently, we tested two alternative silvicultural treatments aimed at improving balsam fir establishment on Anticosti Island (Québec, Canada). In 1998 and 1999, we set up shelterwood seed cutting using three harvest intensities (0, 25 and 40% of basal area) and strip clearcutting with scarification using three different strip widths (15, 30 and 45 m), both with fenced and unfenced regeneration plots, in balsam fir stands. After 8 years, shelterwood seed cutting did not allow the establishment of new balsam fir seedlings, nor the development of unbrowsed balsam fir seedlings. In the strip clearcutting, deer browsing suppressed growth of palatable species in all strip widths. This favoured the development of unpalatable species, especially white spruce (Picea glauca (Moench) Voss). Our study demonstrates that the use of silvicultural treatments alone is unlikely to restore balsam fir regeneration on Anticosti Island, as long as the deer population remains higher than 20 deer/km².     

Pathogenicity of Phytophthora species to Pacific Northwest Conifers

Phytophthora root rot is described for the first time killing sugar pine (Pinus lambertiana) in a seed orchard and four species of true fir (Abies spp.) in a forest nursery. P. cactorum was recovered from true firs and P. megasperma was recovered from sugar pine. P. cryptogea was recovered from sugar pine and true fir but isolates from the two locations differed from each other in pathogenicity and colony appearance. Isolates recovered from these hosts and isolates of 6 Phytophthora species previously recovered from Douglas-fir (Pseudotsuga menziesii) were then tested for pathogenicity on seedlings of 9 Northwest conifers. P. megasperma Group 1, P. cryptogea, and P. cinnamomi were pathogenic to all tree species except western redcedar (Thujaplicata). Western hemlock (Tsuga heterophylla) and true firs were susceptible to most species tested; ponderosa (P. ponderosa) and sugar pines were damaged only by P. cryptogea and P. cinnamomi; western redcedar was resistant to all isolates.     

Short-term browsing by roe deer has little effect on survival and growth of sessile oak seedlings

We studied the effect of roe deer (Capreolus capreolus) browsing on the growth and mortality of sessile oak (Quercus petraea) seedlings. First, we measured the height and basal diameter, and counted the number of shoots of the 849 seedlings of an experimental plantation, 60 of which were individually protected by fencing. We then introduced two tame roe deer females every day for three weeks from late May to mid-June. Measurements and counts were repeated after roe deer removal, then once again the following winter. We found no significant effect of browsing on mortality, total height, or basal diameter of the seedlings. However, the browsed seedlings exhibited a lower number of shoots despite regrowth observed after roe deer removal. We propose to use the tolerance of oak to deer browsing to promote natural and inexpensive methods, to protect oak during regeneration, such as the spatial arrangement of plants or the use of slash or surrounding vegetation in favourable environments.     

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.     

Midcanopy growth following thinning in young-growth conifer forests on the Olympic Peninsula western Washington

Midcanopy layers are essential structures in “old-growth” forests on the Olympic Peninsula. Little is known about which stand and tree factors influence the ability of midcanopy trees in young-growth forests to respond to release; however, this information is important to managers interested in accelerating development of late-successional structural characteristics. We examined basal area growth response of midcanopy trees following variable-density thinning in an effort to determine the effect of thinning and local environment on the release of western hemlock (Tsuga heterophylla (Raf.) Sarg.) and western redcedar (Thuja plicata ex. D. Don) on the Olympic Peninsula in western Washington. Release was measured as the difference between average annual basal area growth over the 5-year prior to thinning and the 3-to-6 year period following thinning. Results indicate that while growth rates were similar prior to thinning (5.4 cm² year⁻¹in both thinned and unthinned patches) midcanopy trees retained in a uniformly thinned matrix grew significantly more (8.0 cm² year⁻¹) than those in unthinned patches (5.4 cm² year⁻¹) for western hemlock and for western redcedar. Crown fullness and crown crowding affected the release of western hemlock in the thinned matrix. Initial tree size, relative age, local crowding and measures of crown size and vigor affected the release of western redcedar in the thinned matrix. Our results indicate that midcanopy western hemlock and western redcedar retain the ability to respond rapidly with increased growth when overstory competition is reduced and the magnitude of response is related to neighborhood variables (intracohort competition, overstory competition, and tree vigor), thus suggest that variable-density thinning can be an effective tool to create variability in the growth of midcanopy trees in young-growth stands. We expect that this rapid response will produce even greater variability over time.     

Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

Effects of browsing on recruitment and mortality of European aspen (Populus tremula L.)

Abstract

High browsing pressure may reduce or halt regeneration of browsing-prone tree species. In this study, exclosures were used to quantify the effects of browsing on vital demographic parameters in European aspen (Populus tremula L.) stands. The experiment focused on stands located in managed forest landscape with densities of 0.85 moose (Alces alces L.)km^?2. Other herbivores such as mountain hare (Lepus timidus L.) were present at densities common for this region and roe deer (Capreolus capreolus L.) at low densities. Recruitment and mortality of aspen ramets were recorded before and after the growing season, and over four consecutive years. Seven to 19% of the ramets within browsing reach were browsed annually. No differences were found in recruitment or mortality attributable to browsing per se. Instead, there were large among-stand variability in vital population parameters unrelated to browsing incidence. In general, aspen stands with high ramet density showed higher recruitment rates than stands with low ramet density. There was substantial variability in self-replacement capability unrelated to ecological factors. The results suggest that more attention should be given to finding and managing stands with high capacity for suckering, as these stands may have a disproportionate impact on aspen recruitment at the landscape scale.     

Effects of wounding and fungal infection with Armillaria ostoyae in three conifer species. II. Host response to the pathogen

Structural responses in the bark and wood were described following penetration by Armillaria ostoyae in the roots of 20‐ to 30‐year‐old Douglas‐fir, western hemlock and western redcedar trees. Tissue necrosis presumably caused by fungal exudates was commonly observed at inoculum contact. In Douglas‐fir and western hemlock, A. ostoyae interfered with the initiation of active defence mechanisms involving the development of a lignified zone of impervious tissue (IT), necrophylactic periderm (NP) formation and compartmentalization of infected woody tissue. Breaching of IT and NP barriers was frequent, particularly around the clusters of sclereid cells in western hemlock. In western redcedar, the IT zone was inconspicuous. Induced rhytidome formation occurred in western redcedar either simultaneously with or after completion of NP development. The formation of this tissue facilitated en masse sloughing of infected tissue from the surface of roots. In western redcedar, traumatic phloem resin ducts formed in tangential bands surrounding the margin of expanded lesions. Effective compartmentalization in western redcedar was achieved by a barrier zone comprised of a higher‐than‐average number of axial parenchyma that accumulated polyphenolic deposits. A combination of host‐mediated defence mechanisms in western redcedar resulted in a significantly higher frequency of effective resistance reactions than in western hemlock or Douglas‐fir.     

Winter Browsing by Large Herbivores on Planted Deciduous Seedlings in Southern Sweden

Deciduous seedlings are valuable for the forest industry and, at the same time, often palatable for large herbivores. Therefore, browsing on seedlings of seven deciduous tree species and on Picea abies(L.) Karst., as a reference species, was tested in a field experiment with the species presented at feeding stations. Twenty stations, each with one seedling per species, were established in each of 15 clear-cuts in southern Sweden. The planting took place in October. Depending on species, between 16 and 85% of the seedlings were browsed after one winter. The descending order, based on number of browsed seedlings, was Quercus robur L . > Alnus glutinosa (L.) Gaertn.> Fagus silvatica L.> Tilia cordata Mill.> Prunus avium L.> Betula pendula Roth> Picea abies (L.) Karst. > Fraxinus excelsior L. On 83% (range 45-100%) of the browsed seedlings the leader was damaged. On average, browsing reduced seedling hight by 32% (range 12-47%). Within species, initially taller seedlings were more often selected than shorter ones.     

Mammalian browsing damage in the Mt. Cole State forest,southeastern Australia: analysis of browsing patterns,spatial relationships and browse selection

Browsing by mammals can have a substantial impact on the survival, growth and form of regenerating forest trees. This study investigated the severity and distribution of mammalian browsing damage, and the degree to which eucalypt seedlings (Eucalyptus spp.) were selected for browsing on three regenerating native forest coupes in southeastern Australia. Damage was greatest 6 months after eucalypt germination and at this time, both messmate (Eucalyptus obliqua) and blue gum (E. globulus) seedlings were browsed at similar intensity. Nine and 12 months after germination, messmate was browsed more heavily than blue gum. Consistent relationships between browsing damage and both distance from the forest edge and wallaby density were not observed. Analysis of use and availability data for messmate, blue gum and manna gum (E. viminalis) seedlings showed that messmate was selected for browsing more than expected by chance while manna gum was browsed less than expected. Mammalian browsing affects a substantial portion of Victorias native forest estate, and could alter the species composition of future forests. Research into the ecology of browsing species and the development of non-lethal browsing reduction strategies is seen as a priority for the management of mammalian browsing in Victoria.     

Browsing by large herbivores on Scots pine (Pinus sylvestris) seedlings in mixture with ash (Fraxinus excelsior) or silver birch (Betula pendula)

Abstract

Food selection by large herbivores occurs at a hierarchy of scales, for example landscape, patch or plant. Several hypotheses regarding food plant selection on patch or plant level have been developed. In this cafeteria-type design field experiment, conducted during one winter immediately after planting, we tested the effect of species mixture on browsing by large herbivores (mainly roe deer) on Scots pine seedlings in mixture with seedlings of ash (highly preferred) or silver birch (less preferred). Browsing on Scots pine was not affected by species mixture, neither in terms of the number of browsed pines nor browsing intensity. Instead, browsed biomass was positively and significantly correlated to the total biomass available for browsing. Also, there were differences due to species, with ash being most browsed (44.6%), followed by Scots pine (18.9%) and silver birch (11.6%). Browsed biomass per browsed seedling, however, was largest for Scots pine. In addition, browsed seedlings were initially taller compared to unbrowsed seedlings for all species. The main management implication in this study is that the species mixture did not influence large herbivore browsing on Scots pine seedlings. Hence, removing or discouraging more (or less) attractive browse species in early stages of pine regeneration activities seems unnecessary from the point of large herbivore browsing.     

Two consecutive yearly applications of orthene medicaps increase protection of grand fir against Western Spruce budworm

A random sample of Douglas-fir, Pseudotsuga menziesii var. glauca (Beissn.) Franco, and grand fir, Abies grandis (Dougl.), in Idaho in the United States, treated with Orthene Medicaps (acephate) in 1979 was treated again in 1980. One year after the second treatment in 1980, twice-treated (treated in 1979 and 1980) and once-treated (treated only in 1979) grand fir had significantly fewer western spruce budworm larvae per 100 buds than did control trees, and twice-treated grand fir had significantly fewer larvae than did once-treated. Defoliation of new shoots on twice-treated and once-treated grand fir was significantly less than that on control trees; defoliation did not differ significantly between twice-treated and once-treated trees for either species. Residues of acephate and methamidophos, its metabolite, monitored in midcrown foliage were detected 1 year after treatment.     

Effects of alternate types of microsite shade on survival of planted Douglas-fir in southwest Oregon

Five-year survival of 2-0 bareroot Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] seedlings was increased by three types of shading (cardboard shadecards placed to the south or east of seedlings, and bottomless styrofoam cups inverted around the seedling base) on two south-facing sites in southwest Oregon. On the drier site, seedlings survived well without shading (89% unshaded, 98% shaded), but on the wetter site, where seedlings were stressed more, shading was more beneficial (62% unshaded, 89% shaded). Shading did not affect growth. Seedlings grew more in 5 years on the drier than the wetter site, likely as a result of better handling and planting practices, less browsing by deer, and better weed control.     

Changes in spatial patterns of sika deer distribution and herbivory of planted seedlings: a comparison before and after deer population control by culling

Sika deer (Cervus nippon) sometimes cause extensive damage to planted tree seedlings. To evaluate the effects of culling on the spatial distribution patterns of sika deer and browse damage to planted seedlings, we compared the data collected before and after experimental culling in a cool, temperate, mixed forest on Kyushu Island, Japan. Experimental culling, conducted in an area of 1 km², removed five, four and two sika deer in April, June and October 2011, respectively. During the year before culling, the spatial pattern of the number of sika deer caught on camera corresponded to the predicted sika deer density. Sika deer immediately browsed planted seedlings after the initial planting. The cumulative number of browsed seedlings increased over time, especially in winter. The spatial pattern of the cumulative number of sika deer caught on camera corresponded to that of browsed seedlings at the year’s end. During the year when culling was conducted, the number of sika deer caught on camera decreased around the center of the study site where the culling was conducted and the number of browsed seedlings decreased. During the year following culling, the cumulative number of browsed seedlings was very similar to that in the year before the culling, while the same low number of sika deer was caught on camera. These results indicate that the effects of deer culling resulted in decreased levels of sika deer appearance and browse damage for more than 1 year and for several months, respectively.