Impacts of non-native plant and animal invaders on gap regeneration in a protected boreal forest

In balsam fir (Abies balsamea)-dominated boreal forests of Gros Morne National Park, Newfoundland (Canada), non-native Cirsium arvense (Canada thistle) has invaded forest gaps. Its management is complicated by the lack of viable control techniques and an overarching issue of gap regeneration failure attributed to browsing by non-native moose (Alces alces). This study identifies the impacts of thistle invasion on balsam fir regeneration and explores protocols to re-establish fir in gaps invaded by thistle and moose. Fir seeds were planted into ten gaps (five natural; five anthropogenic) and the emergence, growth, herbivory damage, and survival of fir was determined for 2 years amongst five treatments (n = 50 plots; 32 seeds/plot): (1) thistle monocultures in gaps; (2) where aboveground thistle biomass was removed; (3) where above- and below-ground thistle biomass was removed; (4) non-invaded areas in gaps; and, (5) adjacent uninvaded forest edges. In addition, 432 fir seedlings (aged 15 months) were transplanted into four forest gaps within the above treatments and followed for 1 year. Results indicate that invasion of C. arvense negatively affects fir emergence and early survival, and may further contribute to continued balsam fir regeneration failure independent of future moose densities. However, older fir seedlings transplanted into thistle monocultures experienced a positive facilitative effect due to the protection thistle provided against small mammal herbivory. Restoration actions that combine moose density reductions with the planting of fir seedlings offers the most viable long-term strategy to re-establish the native forest canopy in thistle-invaded gaps and would likely lead to the eventual decline of shade-intolerant C. arvense.     

Effects of position, understorey vegetation and coarse woody debris on tree regeneration in two environmentally contrasting forests of north-western Patagonia: a manipulative approach

Aim To investigate the differential effects of position within gaps, coarse woody debris and understorey cover on tree seedling survival in canopy gaps in two old‐growth Nothofagus pumilio (Poepp. & Endl.) Krasser forests and the response of this species to gaps in two forests located at opposite extremes of a steep rainfall gradient. Location Nahuel Huapi National Park, at 41° S in north‐western Patagonia, Argentina. Methods In both study sites, seedlings were transplanted to experimental plots in gaps in three different positions, with two types of substrate (coarse woody debris or forest floor), and with and without removal of understorey vegetation. Survival of seedlings was monitored during two growing seasons. Soil moisture and direct solar radiation were measured once in mid‐summer. Seedling aerial biomass was estimated at the end of the experiment. Results Mid‐summer soil water potential was lowest in the centre of gaps, in plots where the understorey had been removed, and highest at the northern edges of gaps. Direct incoming radiation was highest in gap centres and southern edges, and lowest at northern edges. Seedling mortality was highest in gap centres, in both sites. Coarse woody debris had a positive effect on seedling survival during summer in the mesic forest and during winter in the xeric forest. The removal of understorey cover had negative effects in gap centres during summer. Seedling final aerial biomass was positively affected by understorey removal and by soil substrate in both sites. In the dry forest gaps, seedling growth was highest in northern edges, whereas it was highest in gap centres in the mesic forest. Overall growth was positively related to survival in the xeric forest, and negatively related in the mesic forest. Main conclusions Survival and growth were facilitated by the shade of gap‐surrounding trees only in the xeric forest. Understorey vegetation of both forests facilitated seedling survival in exposed microsites but competed with seedling growth. Nurse logs were an important substrate for seedling establishment in both forests; however, causes of this pattern differed between forests. Water availability positively controls seedling survival and growth in the xeric forest while in the mesic forest, survival and growth are differentially controlled by water and light availability, respectively. These two contrasting old‐growth forests, separated by a relatively short distance along a steep rainfall gradient, had different yet unexpected microenvironmental controls on N. pumilio seedling survival and growth. These results underscore the importance of defining microscale limiting factors of tree recruitment in the context of large‐scale spatial variation in resources.     

The impact of simulated moose densities on conifer aphids along a productivity gradient

Large herbivores have a significant impact on boreal forest ecosystems. The modification of resources through their feeding induces changes in trophic dynamics and affects the direction of interactions in a community. Moose Alces alces may decrease the available plant biomass for herbivorous insects on one of their main winter forage species in Fennoscandia, Scots pine Pinus sylvestris, and indirectly alter the abundance of invertebrates through exploitative competition. Moose browsing can also induce chemical, morphological, and phenological changes in plants, changing their nutritive value to insect herbivores such as aphids. Habitat productivity may further modify the responses of aphids to moose browsing. We studied the responses of the gray pine aphid Schizolachnus pineti to different moose densities, and their relations to habitat productivity by sampling pine branches and measuring the number of aphids on pine needles. The experimental setup consisted of 8 exclosures along a productivity gradient, where the feeding, defecation, and urination of 4 densities of moose had been simulated for 7 yr. We here show that high levels of simulated browsing decrease the amount of gray pine aphids in areas with high productivity. In areas with low productivity, however, simulated browsing had no such effect. Habitat productivity should therefore be considered as an important factor that may determine the strength of an areas buffering capacity to high moose densities. Low resource environments appear to be favourable to specialist conifer aphids on pines under high browsing pressure, but the performance of generalist insect herbivores might be lowered.     

Smaller and more numerous harvesting gaps emulate natural forest disturbances: a biodiversity test case using rove beetles (Coleoptera,Staphylinidae)

Aim To evaluate changes in the abundance, species richness and community composition of rove beetles (Coleoptera, Staphylinidae) in response to three configurations of experimental gap cuts and to the effects of ground scarification in early succession yellow birch‐dominated boreal forest. In each experimental treatment, total forest removed was held constant (35% removal by partial cutting with a concomitant decrease in gap size) but the total number of gaps was increased (two, four and eight gaps, respectively), resulting in an experimental increase in the total amount of ‘edge’ within each stand. Location Early succession yellow birch‐dominated forests, Quebec, Canada. Methods Pitfall traps, ANOVA, MIXED procedure in sas ®, post hoc Tukey's adjustment, rarefaction estimates, sum‐of‐squares and distance‐based multivariate regression trees (ssMRT, dbMRT). Results Estimates of species richness using rarefaction were highest in clearcut and two‐gap treatments, decreased in smaller and more numerous gaps and were significantly higher in scarified areas than in unscarified areas. ANOVA indicated a significant impact of harvesting on the overall standardized catch. Post hoc Tukey's tests indicated that the total catch of all rove beetles was significantly higher in uncut forests than in the treated areas. Both sum‐of‐squares and distance‐based multivariate regression trees indicated that community structure of rove beetles differed among treatments. Assemblages were grouped into (a) control plots, (b) four‐ and eight‐gap treatments and (c) two‐gap and clearcut treatments. Main conclusions Rove beetle composition responded significantly to increasing gap size. Composition among intermediate and small‐sized gap treatments (four‐ and eight‐gap treatments) was more similar to uncut control forests than were larger gap treatments (two‐gap) and clearcuts. Effects of scarification were nested within the harvested treatments. When the total area of forest removed is held constant, smaller, more numerous gaps are more similar to uncut control stands than to larger gaps and falls more closely within the natural forest heterogeneity.     

Gap disturbances in northern old‐growth forests of British Columbia,Canada

Abstract. We characterized the abundance, size and spatial patterning of canopy gaps, as well as gap‐forming processes and light availability in boreal, sub‐boreal, northern temperate and subalpine old‐growth forests of northwestern British Columbia. The proportion of area in canopy gaps ranged from 32% in northern temperate forests to 73% in subalpine forests. Evenly distributed developmental gaps were dominant but permanent openings created by edaphic components and by shrub communities were also common, particularly in subboreal forests. Abundant gaps, large gap sizes, high numbers of gap makers per gap and frequent gap expansion events suggest that gaps have long tenure in these forests. Snapped stems and standing dead mortality were the most common modes of mortality in all forest types resulting in little forest floor disturbance, creating few germination sites for seedling establishment. We found high mean light levels (16–27% full sun) and little difference between non‐gap and gap light environments. Our results suggest that gap dynamics in these forests differ fundamentally from those in temperate and tropical forest ecosystems.     

Conservation of wildlife populations: factoring in incremental disturbance

Progressive anthropogenic disturbance can alter ecosystem organization potentially causing shifts from one stable state to another. This potential for ecosystem shifts must be considered when establishing targets and objectives for conservation. We ask whether a predator–prey system response to incremental anthropogenic disturbance might shift along a disturbance gradient and, if it does, whether any disturbance thresholds are evident for this system. Development of linear corridors in forested areas increases wolf predation effectiveness, while high density of development provides a safe‐haven for their prey. If wolves limit moose population growth, then wolves and moose should respond inversely to land cover disturbance. Using general linear model analysis, we test how the rate of change in moose (Alces alces) density and wolf (Canis lupus) harvest density are influenced by the rate of change in land cover and proportion of land cover disturbed within a 300,000 km² area in the boreal forest of Alberta, Canada. Using logistic regression, we test how the direction of change in moose density is influenced by measures of land cover change. In response to incremental land cover disturbance, moose declines occurred where <43% of land cover was disturbed; in such landscapes, there were high rates of increase in linear disturbance and wolf density increased. By contrast, moose increases occurred where >43% of land cover was disturbed and wolf density declined. Wolves and moose appeared to respond inversely to incremental disturbance with the balance between moose decline and wolf increase shifting at about 43% of land cover disturbed. Conservation decisions require quantification of disturbance rates and their relationships to predator–prey systems because ecosystem responses to anthropogenic disturbance shift across disturbance gradients.     

Impact of moose population density on the production and composition of litter in boreal forests

Recent studies of ungulates have revealed that selective foraging seems to be an important mechanism by which they can affect the structure and species composition of the plant community, and thus quantity (dry mass) and quality (chemical composition) of litter available for decomposers. Such changes in litter production may be especially important in N-limited systems like boreal forests. We chose moose ( Alces alces ) as study species to investigate this mechanism. Moose browse mainly in the tree and shrub layers year round, and because of their wide distribution and often high population densities, they can have a significant effect on litter production of trees and shrubs in Swedish boreal forests. The effects of herbivores may also vary along productivity gradients. We therefore simulated browsing and urine and fecal deposition corresponding to 4 different moose densities in exclosures along a pre-existing forest productivity gradient. Both litter quantity (g dry mass per m² and year) and contributions of C and N (g dry mass per m² and year) decreased with increasing level of simulated moose density. High moose densities over extended time can therefore reduce N contributions to soil and therefore eventually reduce site productivity in Swedish boreal forests. This effect of moose was mainly a result of decreased litter quantity, because contradictory to studies from North America, litter quality (C:N ratio and N contribution per mass unit of litter) was not affected by level of simulated moose density.     

Small gap dynamics in the southern boreal forest of eastern Canada: Do canopy gaps influence stand development?

Question: To what extent do small‐scale disturbances in the forest canopy, created by natural disturbance agents, affect stand development? Doubts exist as to whether small canopy openings have any real effect on the understory tree recruitment, especially in boreal forests. Location: Conifer and mixed stands in the Gaspesian region in eastern Québec. The main natural disturbance agents are recurring outbreaks of Choristoneura fumiferana (eastern spruce budworm) and winds. Methods: Linear transects in 27 sites were used to describe the gap (< 0.1 ha) regime parameters, including gap fraction, gap size and change in disturbance severity through time. Three stand types were distinguished, based on a gradient of abundance of tree host species for the eastern spruce budworm. The impact of gaps was evaluated on the basis of changes in the number, the period of recruitment, and the composition of tree saplings present within gap areas. Changes were measured along the gap size gradient, and according to the pattern of recent budworm epidemics. Results: The gap fraction is highly variable (18%‐64%) and is on average relatively high (42%). Gap sizes have a positively skewed distribution. In most cases the growth rate among gap filling saplings increased sufficiently to date disturbance events. The composition and the structure of understory trees were affected by gap formation. The number of shade‐intolerant tree species did increase during or following periods of particularly severe canopy disturbances. However, the establishment or survival of shade intolerant species was not restricted to larger gaps or more intensely disturbed periods. Conclusions: In sub‐boreal forests of Eastern Canada, small scale disturbances in the tree canopy influence stand regeneration dynamics, but not to the extent that parameters such as sapling composition and recruitment patterns depend on gap regime characteristics.     

Artificial canopy gaps accelerate restoration within an exotic Pinus radiata plantation

We created small‐scale artificial canopy gaps to accelerate the growth of mature indigenous forest canopy species for restoration of an 18‐year‐old exotic Pinus radiata plantation forest, in the Marlborough Sounds, New Zealand. Small and large circular gaps were formed by felling. Seedlings of two indigenous forest canopy species, Podocarpus totara (Podocarpaceae) and Beilschmiedia tawa (Lauraceae), were planted within artificial gaps and undisturbed plantation canopy. Seedling height growth, mortality, and occurrence of animal browse were monitored at approximately 6‐month intervals over 17 months. Both P. totara and B. tawa differed significantly in height growth and in animal browse occurrence among artificial gap treatments. Growth of the light‐demanding P. totara was better under large canopy gaps, whereas growth of the shade‐tolerant B. tawa increased under gaps of any size but was most consistent under small gaps. For P. totara, any significant restoration benefit of gap formation on height growth was lost when browsed seedlings were taken into account. Animal browse significantly limited B. tawa height growth in large but not in small gaps. Small‐scale canopy gap creation is an effective method of modifying light transmission to the plantation understorey and accelerating seedling growth rates. Canopy gap size can be used to optimize understorey illumination according to species‐specific light requirements. The increased occurrence of animal browse in gaps requires consideration. Artificial canopy gaps within planted monocultures create structural heterogeneity that would otherwise take an extended period of time to develop. These results further support the role of plantations as indigenous forest restoration sites.     

The role of moose <Emphasis Type="Italic">Alces alces</Emphasis> L. in boreal forest – the effect on ground beetles (Coleoptera,Carabidae) abundance and diversity

The indirect effect of moose Alces alces browsing on ground beetle's abundance and diversity was investigated by pitfall trapping in a mixed coniferous forest in Vestfold County (59°19′ N, 9°50′ E, Norway), during the summer of 2002. Three areas with different browsing pressures, ranging from non- to medium- and heavily browsed were chosen and dry weight of bilberry Vaccinium myrtillus and humidity at ground level were measured in the three locations. As predicted, the gradient analyses showed that browsing by moose influenced the composition of carabid fauna, and that browsing intensity and humidity covaried with the most important gradient in carabid species composition found across the three locations. Species that live in light stands with rather dry soil, were more often captured in the browsed areas, whereas shade tolerant and hygrophilous species were more abundant in the non-browsed area. The carabid abundance increased significantly with increasing browsing pressure. According to our predictions, the diversity at trap level (α-diversity) was higher in the highly browsed area. Conversely, species turnover (β-diversity) decreased with browsing intensity. On the other hand, the rarefaction analysis showed that the regional species richness (γ-diversity) was considerably higher in the medium browsed area than in the heavily browsed one, which is consistent with the intermediate disturbance hypothesis. This study indicates that moose, by reducing the bilberry that constitutes the field layer in summer, affect carabid species composition and might be capable to reshape the whole ecosystem in our study area by a cascade effect.     

Linking radial growth patterns and moderate‐severity disturbance dynamics in boreal old‐growth forests driven by recurrent insect outbreaks: A tale of opportunities,successes, and failures

In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests.     

Feathermoss seedbeds facilitate black spruce seedling recruitment in the forest–tundra ecotone (Labrador,Canada)

As climate warms, conifers are expected to expand their ranges into alpine tundra where ecological factors such as seedbed availability, and post‐dispersal seed and seedling predation may control local recruitment. Seedbed composition may influence microhabitat, nutrients, physical structure, and predation level and, therefore, affect the success of conifer recruitment, thereby providing the template for future expansion. In the boreal forest, seedbed–seedling competition dominates such that seedbed removal increases black spruce recruitment. In the harsher climate of the Mealy Mountains boreal forest–tundra ecotone (Labrador, Canada) the Stress gradient hypothesis (SGH) predicts that facilitation may dominate seedbed–seedling interactions. This study investigated potential mechanisms of seedbed facilitation (temperature, water, nutrients, physical protection) in three conifer seedbeds (Pleurozium schreberi, Cladonia spp., bare soil) and examined whether seed predation and/or seedling herbivory varied among seedbeds over three years. Seed emergence was low overall (< 10% on all treatments), but highest on Pleurozium (6.3%), followed by bare ground (4.6%) and Cladonia (0.3%). Facilitation was observed between Pleurozium and black spruce as seedling height increase (31%) and survival (55%) were highest; herbivory, seed predation and overwinter mortality were lowest compared to both Cladonia and bare ground seedbeds. Unlike in the closed canopy boreal forest, seedlings recruited poorly on bare soil as seedling height increase and survival were 20.5% and 26%. Temperature and water availability were similar across seedbeds, while nutrient availability was higher on Pleurozium. The physical structure of Pleurozium likely protects first to third‐year seedlings from temperature extremes and predators. As climate warms and seed availability increases, Pleurozium may facilitate black spruce recruitment and treeline expansion.     

Leaf litter arthropod responses to tropical forest restoration

Soil and litter arthropods represent a large proportion of tropical biodiversity and perform important ecosystem functions, but little is known about the efficacy of different tropical forest restoration strategies in facilitating their recovery in degraded habitats. We sampled arthropods in four 7‐ to 8‐year‐old restoration treatments and in nearby reference forests. Sampling was conducted during the wet and dry seasons using extractions from litter and pitfall samples. Restoration treatments were replicated in 50 × 50‐m plots in four former pasture sites in southern Costa Rica: plantation – trees planted throughout the plot; applied nucleation/islands – trees planted in patches of different sizes; and natural regeneration – no tree planting. Arthropod abundance, measures of richness and diversity, and a number of functional groups were greater in the island treatment than in natural regeneration or plantation treatments and, in many cases, were similar to reference forest. Litter and pitfall morphospecies and functional group composition in all three restoration treatments were significantly different than reference sites, but island and plantation treatments showed more recovery than natural regeneration. Abundance and functional group diversity showed a much greater degree of recovery than community composition. Synthesis and applications: The less resource‐intensive restoration strategy of planting tree islands was more effective than tree plantations in restoring arthropod abundance, richness, and functional diversity. None of the restoration strategies, however, resulted in similar community composition as reference forest after 8 years of recovery, highlighting the slow rate of recovery of arthropod communities after disturbance, and underscoring the importance of conservation of remnant forests in fragmented landscapes.     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Regeneration response and seedling bank dynamics on a Dendroctonus rufipennis‐killed Picea engelmannii landscape

Question: How does regeneration response to a host‐specific, high‐severity, infrequent Dendroctonus rufipennis outbreak differ from our conceptualization of high‐severity, infrequent/low‐severity, frequent disturbance regimes in Picea engelmannii‐Abies lasiocarpa communities? Location: Southern Utah, USA. Methods: One hundred and seven plots across a high‐elevation P. engelmannii forest were sampled to reconstruct pre‐outbreak overstory and seedling bank densities, and calculate their associated metrics of diversity. Decade of establishment by seedling bank trees indicated “chronic” and “pulse” regenerators. Results: The post‐outbreak overstory and seedling bank were dominated by A. lasiocarpa. Although Pinus flexilis, Pinus ponderosa, Picea pungens, and Psuedotsuga menziesii were present in the overstory, they were virtually absent in the seedling bank. Seedling bank recruitment of A. lasiocarpa and P. engelmannii has been occurring chronically for at least the last ～205 and ～152 years, respectively. A pulse response of seedling bank Populus tremuloides was apparent; however, results were complicated by intense ungulate browsing. Conclusions: Despite some similarities to the high‐severity, infrequent/low‐severity, frequent conceptualization of regeneration response to disturbance, the high‐severity D. rufipennis outbreak is best described by explicitly considering host specificity and severity. Although, the outbreak simultaneously promoted both a pulse of P. tremuloides and a release of chronically regenerated A. lasiocarpa, the P. tremuloides response was generally masked by ungulate browsing, and the regeneration response came overwhelmingly from the A. lasiocarpa seedling bank. In this landscape, once dominated by P. engelmannii, the chronically regenerating seedling bank, typically thought to take advantage of canopy gaps associated with low‐severity disturbances, is poised to dominate forest reorganization in response to the host‐specific outbreak.     

Impact of simulated moose densities on abundance and richness of vegetation,herbivorous and predatory arthropods along a productivity gradient

Large herbivores can affect vegetation structure and species composition as well as material and energy flows in the ecosystem through their selective feeding, defecation, urination and trampling. These changes have a large potential to indirectly affect other trophic levels, but the mechanisms are poorly known. We studied the impacts of moose Alces alces browsing along a gradient of site productivity by experimentally simulating four different moose densities. Here we show that moose can affect the richness and abundance of three trophic levels in Swedish boreal forests through complex direct and indirect impacts, but in qualitatively different ways depending on how the physical habitat or food resources of a trophic level are affected. Vegetation richness had a hump‐shaped (unimodal) response to increased moose density. Leaf litter production decreased when browsing increased, which in turn depressed the abundance of flying prey for spiders. Consequently, spider abundance and richness declined monotonically. The responses of spider richness to moose density were further conditioned by site productivity: the response was positive at productive and negative at unproductive sites. In contrast, herbivorous Hemiptera were not affected by moose, most likely because the abundance of their food plants was not affected. The highest simulated moose density had an impact on all variables responding to moose even after a few years of treatment and can be considered as overabundance. We also show that the impacts of low or moderate moose density can be positive to some of the organisms negatively affected by high density. The level of herbivore population density that leads to substantial community impacts also depends on site factors, such as productivity.     

Shift of Conifer Boreal Forest to Lichen–Heath Parkland Caused by Successive Stand Disturbances

In the southern boreal forest (Québec, Canada), tree harvesting is a major disturbance affecting the dominant black spruce (Picea mariana) stands already suffering from naturally recurrent insect and fire disturbances. Although recovery of the spruce forest after an insect infestation or a fire is possible under current site conditions, it is less likely when both types of disturbance occur during a short period of time. The addition of yet another disturbance, such as tree harvesting, can thus have catastrophic consequences. We analyzed the impact of three successive disturbances—tree harvesting, insect infestation, and fire—on the regeneration of boreal spruce–moss forests within a period of approximately 50 years. The spruce forests were harvested in the 1940s and the 1950s. Recovery from the logging consisted of advance regeneration (spruce layers less than 1 m high that were left intact during clear-cuts), which was burned in 1991. The vegetation cover (mostly heath and lichen species) and soil conditions (acidic, nutrient-poor podzolic soils developed from coarse materials) of the postfire sites that we studied were similar. Stand structure and tree regeneration were documented from large quadrats (0.25 ha) using age, size, and tree ring data from postlogged and postfire spruce. At an early stage of development, the growing advance regeneration was damaged by insect defoliators in the late 1970s and the mid-1980s, and several trees died a few years before the 1991 fire. The successive disturbances considerably reduced the number of seed-bearers, leading to the collapse of postfire regeneration and a shift to parkland. Through a successional trajectory far from the expected trend for boreal forests influenced by single disturbance, the shift resulted in the formation of divergent plant communities. The development of divergent communities at the landscape scale is generally overlooked due to their small size. They indicate, however, the weak resilience of boreal forests faced with cascading perturbations, which are likely to increase in intensively logged areas.     

Short-term Effects of Harvest Technique and Mechanical Site Preparation on Arthropod Communities in Jack Pine Plantations

Arthropods play a key role in the functioning of forest ecosystems and contribute to biological diversity. However, the influence of current silvicultural practices on arthropod communities is little known in jack pine (Pinus banksiana) forests, a forest type comprising a major portion of the Canadian boreal forest. In this study, the effects of silvicultural treatments on arthropod communities were compared to identify those treatments that minimize ecological impacts on arthropods. The influence of harvesting techniques and mechanical site preparations on insect family richness and abundance of arthropods (total, by orders and by trophic groups) was examined in young (three-year-old) jack pine plantations of northern Ontario. Each of the following treatments were conducted in three plots: (1) tree length harvest and trenching; (2) full tree harvest and trenching; (3) full tree harvest and blading; and (4) full tree harvest and no site preparation. Arthropods were collected using sweepnets and pitfall traps over two years. Blading significantly reduced insect family richness, the total abundance of arthropods, abundance of Orthoptera, Heteroptera, Hymenoptera, Diptera, insect larvae, and plant feeders when compared to the other treatments. The use of either full tree or tree length harvesting had similar short-term effects on family richness and the abundance of arthropods. Arthropod diversity declined with increasing post-harvest site disturbance. These results suggest that arthropod communities in the understory and on the ground are reduced most on sites mechanically prepared by blading, but are similar under conditions immediately following either full tree or tree length harvesting. The implications for regenerating jack pine in the boreal forest are discussed.     

Ungulate herbivory reduces abundance and fluctuations of herbivorous insects in a boreal old-growth forest

Ecological theory predicts the strongest ecosystem effects of herbivory when dominant and ecologically important species are consumed. Bilberry, Vaccinium myrtillus, is such a key plant species, attractive to many other species in the boreal forests, for example ungulate and invertebrate herbivores. Large herbivores may remove substantial biomass and alter plant quality and therefore affect abundance and populations of invertebrate animals sharing the same food plant. We combined experimental exclusion of ungulates with a browsing intensity gradient to investigate the 15-year effect of ungulate (Cervus elaphus and Ovis aries) browsing on bilberry plant size and on bilberry-feeding herbivorous larvae (Lepidoptera and Symphyta), in a Norwegian old growth boreal forest ecosystem. Bilberry ramets in exclosure plots had nearly nine times higher dry mass and three times higher abundance of invertebrates feeding on them than in ungulate-access plots. Sweep-netting data verified these findings as larval numbers were twice as high in exclosure plots. The pattern in the large herbivore effects on bilberry size and abundance of herbivorous larvae were identical along the browsing gradient. Differences in larval abundance between treatments, as indicated by leaf-chewing, increased during the 15-year study period, and the community fluctuations were larger when ungulate herbivores were excluded. The browsing effect was moderated by plant quality as larval densities were lowest on both heavily-browsed and non-browsed plants, and highest on ramets that had 50–74% of annual shoots browsed. Our study supports previous findings in that bilberry is relatively disturbance tolerant and may recover quickly, but that ungulates may compete with herbivorous larvae for food biomass. Additionally, our results strongly indicates that population insect community peaks and fluctuations are dampened by ungulate consumption. Our findings add to the understanding on how ungulates may structure forest ecosystems directly and indirectly.     

Response of ground and rove beetles (Coleoptera: Carabidae,Staphylinidae) to operational oil sands mine reclamation in northeastern Alberta,a case study

Species loss caused by anthropogenic disturbance threatens forest ecosystems globally. Until 50 years ago, the major sources of boreal forest disturbance in western Canada were a combination of forest wild fire events, pest insect outbreaks, and forest timber harvesting. However, in the 1960s, when the oil boom started in Alberta, oil and gas development along with oil sands mining quickly became another major forest disturbance agent. In this case study we report the effects of operational oil sands mine reclamation on terrestrial arthropod communities and compare them with nearby burned and mature forest sites as a way to provide a benchmark from which to understand the long-term trajectory of recovery for these groups. During the summer of 2016 over 6700 epigaeic beetles were collected using pitfall traps. A total of 43 species of ground beetles and 118 species of rove beetles were collected. Epigaeic beetle assemblages differed between the reclaimed, burned, and mature forest sites. Partitioning of beta diversity in the reclaimed, burned areas and mature forests indicated that species turnover formed the largest component of diversity. Species richness patterns were similar among sites; however, cluster analysis indicated that epigaeic beetle assemblages were only 20% similar between the reclaimed and natural sites. Although ground beetles of the reclaimed area showed positive spatial autocorrelation among treatments, both ground and rove beetles showed responses to the reclamation treatments. The reclaimed areas were dominated by small- to medium-sized open-habitat eurytopic species, whereas the fire and mature forest sites were dominated by larger forest species. The reclaimed area of this case study constitutes a novel, reconstructed ecosystem that is clearly not equivalent in species assemblage to burnt stands of similar age or to mature forest stands.