Tree mortality and habitat shifts in the regeneration trajectory underneath canopy of an old-growth subalpine forest

To better understand tree regeneration trajectories and the resultant coexistence of Abies with co-dominants, Picea jezoensis var. hondoensis, Tsuga diversifolia and Betula ermanii, in an old-growth subalpine forest, we investigated spatial mortality patterns during the regeneration of Abies mariesii and A. veitchii, which are abundant in the understory reflecting their shade tolerance. Regeneration of these Abies spp. from shaded understory to canopy status is affected by other canopy co-dominants. Snags of understory Abies spp. were common, suggesting that the primary mortality agent is suppression by the overstory. Although live, small Abies trees in the understory were positively associated with a Picea canopy, the long-term survival was reduced among Abies trees close to the canopy, suggesting that shading by large Picea in the overstory negatively affects understory Abies plants. The existence of shade-intolerant canopy co-dominants such as Picea and also Tsuga, which are larger and longer lived than the shade-tolerant Abies, may play an important role in preventing the Abies spp. from competitively displacing these other tree species, which are much rarer in the understory, though common in the canopy. Moreover, in spite of the fact that Betula canopies fostered recruitment and growth of Abies saplings, Abies showed no association with Betula canopy and their survival at later-stage was rather reduced near or beneath Betula canopies at the subsequent understory small tree stage. Based on spatially significant events related to tree death, this study detected such “habitat shifts” in the trajectory of tree regeneration. Accordingly, it can be concluded that careful consideration of the regeneration habitat is required for a fuller understanding of ecological processes in spatially complex old-growth forest systems.     

Relationships between advance oak regeneration and biotic and abiotic factors

Relationships between advance regeneration of four tree species (red maple (Acer rubrum L.), white oak (Quercus alba L.), chestnut oak (Q. montana Willd.) and northern red oak (Q. rubra L.)) and biotic (non-tree vegetation and canopy composition) and abiotic (soil series and topographic variables) factors were investigated in 52, mature mixed-oak stands in the central Appalachians. Aggregate height was used as a composite measure of regeneration abundance. Analyses were carried out separately for two physiographic provinces. Associations with tree regeneration were found for all biotic and abiotic factors both in partial models and full models. Red maple was abundant on most of the sites, but high red maple abundance was commonly associated with wet north-facing slopes with little or no cover of mountain-laurel (Kalmia latifolia L.) and hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore). Regeneration of the three oak species was greatly favored by the abundance of overstory trees of their own kind. White oak regeneration was most abundant on south-facing, gentle, lower slopes with soils in the Buchanan series. Chestnut oak regeneration was more common on south-facing, steep upper slopes with stony soils. There was a positive association between chestnut oak and huckleberry (Gaylussacia baccata (Wangh.) Koch) cover classes. Northern red oak was more abundant on north-facing wet sites with Hazleton soil, and was associated with low occurrence of mountain-laurel and hay-scented fern.     

The mating and oviposition behavior of the invasive emerald ash borer (Agrilus planipennis), with reference to the influence of host tree condition

Emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is an invasive wood-boring beetle first detected in North America in 2002. Relatively little is known about the factors affecting the mating and oviposition behavior of EAB, even though they could have implications for the management of this species. Consequently, we conducted field surveys in Michigan to observe the diurnal behavior of EAB adults, in addition to examining how their behaviors associated with reproduction varied in relation to the size and condition of host trees. We observed most EAB adults between 11:00 and 15:00, and this pattern was the same for both sexes and for all of the specific behaviors examined (feeding, flying, mating, resting, and walking). Regarding host tree size and crown condition, we consistently found that the most EAB eggs, adults, mating pairs, and ovipositing females were observed on intermediately stressed trees with 40–60 % crown reductions (likely resulting from earlier EAB infestations). Additionally, host tree crown condition appeared to be a more important factor than diameter at breast height. Our results therefore provide support to the hypothesis that host tree crown (stressing) condition influences EAB oviposition behavior. Determining which trees are likely to contain the most EAB eggs should help to guide efforts for releasing and promoting the establishment of parasitoids utilized in biologic control. However, future work should attempt to experimentally test the hypothesis that host tree condition (e.g., crown reduction) drives EAB oviposition behavior, and investigate the fitness implications of these host choices.     

Prescribed burning and understory composition in a temperate deciduous forest,Ohio, USA

Since the advent of widespread suppression in the mid-20th century, fire has been relatively rare in deciduous forests of the eastern United States. However, widespread prescribed burning has recently been proposed as a management tool to favor oak (Quercus spp.) regeneration. To examine the potential effects of fire introduction on the understory community, we experimentally burned small plots and simulated aspects of fire at a forested site in southeastern Ohio. Treatments included two burn intensities, litter removal, increased soil pH, and a control. Treatments were arranged in a randomized block design in two landscape positions (dry upland and moist lowland) and two canopy conditions (gap, no gap). Post-fire vegetation was identified to species, and stems were counted 1, 3, and 14 months after burning. Community composition was more strongly affected by fire in upland plots than in lowlands, but was not affected by canopy openness. Both cool and hot burns reduced post-fire seedling emergence of Acer rubrum, a common overstory tree. Hot burns facilitated germination of Vitis spp., Rhus glabra, and Phytolacca americana, species common in disturbed habitats, and increased graminoid abundance. Cool burns and litter removal facilitated germination of Erechtites hieracifolia and Liriodendron tulipifera suggesting that litter removal is the mechanism by which fire favors colonization. These results suggest that fire applied frequently in the Central Hardwoods Region would cause compositional shifts to graminoids and disturbance-adapted forbs by increasing germination from the seed bank. Fire did not favor species with dormant underground buds, as studies in other ecosystems would suggest. Vegetational responses were noticeably weaker in the second year after burning, indicating that a single fire has only a short-term effect.     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.     

Recruitment limitation in forests: Lessons from an unprecedented mountain pine beetle epidemic

Since the mid-1990s the forests of central British Columbia have undergone an unprecedented Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) (MPB) epidemic that has resulted in extensive mortality of canopy lodgepole pine (Pinus contorta var. latifolia Engelm.). This study investigated how seed-source availability, seedbed substrate, overstory structure, and time since MPB attack interact to affect post-MPB seedling recruitment of the dominant tree species of these forests. In addition to post-MPB recruitment, these forests may be regenerated by trees established in the understory prior to MPB disturbance. Accordingly, we examined abundance and patterns of all regeneration less than 130 cm tall. We found post-MPB recruitment was sparse. Subalpine fir (Abies laciocarpa (Hook.) Nutt.) comprised the majority of the post-MPB recruitment. It increased with local parent tree basal area and increased strongly with proximity to a major seed source. This resulted in a patchy distribution for subalpine fir post-MPB regeneration. Lodgepole pine post-MPB recruitment was limited by overstory shading. Recruitment of pine decreased as the total overstory basal area increased. Interior spruce (Picea glauca × engelmannii) post-MPB recruitment was similarly limited by total overstory basal area. Seedbed substrates were uniform and dominated by moss. Substrate type distribution did not change as time since MPB disturbance increased. The overall low post-MPB recruitment observed was likely due to a lack of disturbance to the moss-dominated forest floor. Moss is known to be a poor substrate in northern forests. The distribution of all regeneration less than 130 cm tall showed the same trends as the post-MPB regeneration. We believe the post-MPB seedling recruitment dynamics of these forests was not substantially changed from conditions prior to MPB disturbance. There was no pulse of regeneration up to 10 years post-MPB disturbance. Unless this changes, future stand structure will be dominated by the seedling bank established prior to the MPB epidemic. Subalpine fir dominated the seedling bank (68%) and post-MPB recruitment (94%). This suggests that MPB-disturbed forests are undergoing a substantial shift in landscape-level species composition.     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Tree recruitment in gaps of various size,clearcuts and undisturbed mixed forest of interior British Columbia,Canada

Tree seedling recruitment was monitored after various types of logging in mixed conifer and deciduous forests of northern British Columbia, Canada. Predicting tree seedling recruitment after disturbance is fundamental to understanding forest dynamics and succession and is vital for forest management purposes. Seedling recruitment success in multi-species northern latitude forests varied as a function of mature tree canopy cover, gap size and position in a gap. Recruitment was abundant within canopy gaps across a wide range of gap sizes (20–5000 m²), but recruit numbers dropped off rapidly under the closed forest canopy and in the open conditions of clearcuts. Inside canopy gaps, recruitment was similar by gap position in small gaps (<300 m²) but, in these northern latitude forests, exhibited a trend of increasing density from the sunny north to shady south end of larger gaps. This was true for all tree species regardless of their shade tolerance ranking. There was no evidence of gap partitioning by any of the tree species during the regeneration phase suggesting that adaptation to the subtleties of gap size during early recruitment are not well developed in these tree species. Favorable locations for emergence and early establishment of germinants were less favorable for growth and survival of established seedlings, i.e. the regeneration niches in these forests were discordant. Tree abundance and species diversity appears to be controlled more by differentiation among growth and survival niches than by the regeneration niches. From the perspective of forest management, abundant natural regeneration of all the dominant tree species of these mixed-species forests can be obtained after partial cutting.     

Understory bamboo flowering provides a very narrow light window of opportunity for canopy-tree recruitment in a neotropical forest of Misiones, Argentina

Chusquea ramosissima is a native monocarpic bamboo species growing in subtropical forests of northeastern Argentina, which can dominate gaps and open forests in the region, particularly after human disturbance. This bamboo species started to flower in different areas of northeastern Argentina in 2001, with the flowering peak during 2002 and 2003 and small isolated flowering events still occurring until 2010. We studied the effects of C. ramosissima flowering and die-back on microclimate, litter decomposition, nutrient availability, sapling growth, abundance and regeneration of tree canopy species. We wanted to know how environmental conditions and ecosystem processes change through time after bamboo flowering and if bamboo die-back would favor regeneration of canopy trees. Twenty 50 × 50 m plots of flowering and non-flowering bamboo were permanently marked and vegetation dynamics as well as nutrient cycling and microclimate studies were performed. C. ramosissima die-back enhanced growth and reduced mortality rate of tree saplings during the first year after flowering. Only growth of tree saplings previously established was enhanced by the flowering event and tree-species richness and saplings abundance of canopy trees did not change as expected due to bamboo flowering. The short-term effect of tree saplings growth was likely due to incident solar radiation at the forest floor which doubled in the first year after the bamboo flowering event. Increased light availability at the forest floor simultaneously promoted the growth of other understory plants such as ferns, lianas and Piper spp. that rapidly colonized gaps and intercepted a percentage of the incident solar radiation after the first year, which together with an increased litter layer due to the senescence of the bamboo, may have inhibited establishment of new tree individuals and affected tree growth. Contrary to predictions, soil water, litter decomposition and soil nutrients were not significantly affected by bamboo flowering. Thus, successful tree regeneration in gaps following bamboo flowering appears to be restricted to a very narrow window of increased light availability (i.e., 1 year) before growth of other understory plants and rapid re-colonization of bamboo. Changes in resource availability, and the opportunity for overstory regeneration after bamboo flowering events appears to depend on climatic and community characteristics of the ecosystem where the flowering event occurs and also, on the flowering patterns and their synchronicity.     

Disturbance severity and canopy position control the radial growth response of maple trees (<Emphasis Type="Italic">Acer</Emphasis> spp.) in forests of northwest Ohio impacted by emerald ash borer (<Emphasis Type="Italic">Agrilus planipennis</Emphasis>)

Key message

Radial growth of silver and red maples was investigated across three forests in northwest Ohio following the outbreak of the invasive emerald ash borer. The growth response of maples was driven by an advancement in canopy class and disturbance severity.

Context

Forest disturbances resulting in species-specific diffuse mortality cause shifts in aboveground and belowground competition. This competition may differentially affect non-impacted trees, depending on crown class, disturbance severity, and species-specific responses.

Aims

The purpose of this study is to elucidate the primary drivers of silver and red maple (Acer saccharinum and A. rubrum) growth following emerald ash borer (EAB, Agrilus planipennis)-induced ash tree (Fraxinus spp.) mortality in riparian forests of northwest Ohio.

Methods

Using dendroecological approaches, we analyzed the pattern of radial growth in red and silver maples in conjunction with the EAB outbreak.

Results

This study revealed growth rates of maples increased 72% following EAB arrival and trees advancing in crown class grew 41% faster than those not advancing. The growth response varied by initial crown class, with trees in the intermediate class responding most dramatically. Furthermore, the positive correlation between relative basal area of ash and the radial growth response of maples indicates the important role of disturbance severity in post-disturbance dynamics.

Conclusion

These findings suggest that, although advancement in crown class may allow predictions of “winners” in forest succession post-disturbance, even trees not changing crown class benefit from decreased competition. Results from this study provide a detailed account of radial growth responses in maples following EAB-induced ash mortality and lend insight into the future canopy composition of ash-dominated riparian forests.

     

The role of gaps and tree regeneration in the transition from dense to open black spruce stands

Black spruce forests growing on clay soils in northwestern Quebec change structure from dense even-aged stands to open uneven-aged stands such that almost all forests older than 200 years have an open canopy. These forests become unproductive over time because they are prone to paludification. The main goal of our study was to document the transition between dense and open stands in terms of gap dynamics, with a focus on tree regeneration. Our objective was to determine whether forests remain open due to a lack of regeneration, a lack of growth or both. Nine stands along a 50–250-year-old time since fire gradient were sampled with the line intersect sampling method. Gap fraction increased with stand age and reached a maximum of 77% in the oldest site. In old-growth stands, gaps were interconnected due to the low density of these forests. Most of the gap makers were found with broken stems. Regeneration was dominated by black spruce layers and was relatively abundant (1.71 stems/m²). However, the majority of gap fillers were smaller than 1 m in height in stands of all ages. Instead of a lack of regeneration, the opening of the forests is due to a lack of growth associated with cold and wet organic deposits. Partial harvesting could be implemented on the most productive sites, while management techniques including soil disturbances will be required on low productivity sites to recreate good growth conditions.     

Habitat associations of saproxylic beetles in the southeastern United States: A comparison of forest types,tree species and wood postures

Saproxylic beetles are highly sensitive to forest management practices that reduce the abundance and variety of dead wood. However, this diverse fauna continues to receive little attention in the southeastern United States even though this region supports some of the most diverse, productive and intensively managed forests in North America. In this replicated three-way factorial experiment, we investigated the habitat associations of saproxylic beetles on the coastal plain of South Carolina. The factors of interest were forest type (upland pine-dominated vs. bottomland hardwood), tree species (Quercus nigra L., Pinus taeda L. and Liquidambar styraciflua L.) and wood posture (standing and downed dead wood, i.e., snags and logs). Wood samples were taken at four positions along each log and snag (lower bole, middle bole, upper bole and crown) ∼11 months after the trees were killed and placed in rearing bags to collect emerging beetles. Overall, 33,457 specimens from 52 families and ≥250 species emerged. Based on an analysis of covariance, with surface area and bark coverage as covariates, saproxylic beetle species richness differed significantly between forest types as well as between wood postures. There were no significant interactions. Species richness was significantly higher in the upland pine-dominated stand than the bottomland hardwood forest, possibly due to higher light exposure and temperature in upland forests. Although L. styraciflua yielded more beetle species (152) than either Q. nigra (122) or P. taeda (125), there were no significant differences in species richness among tree species. There were also no relationships evident between relative tree abundance and observed or expected beetle species richness. Significantly more beetle species emerged from logs than from snags. However snags had a distinct fauna including several potential canopy specialists. Our results suggest that conservation practices that retain or create entire snags as opposed to high stumps or logs alone will most greatly benefit saproxylic beetles in southeastern forests.     

六盘山区辽东栎林分结构对各植物层生物量的影响

目的 量化宁夏六盘山区辽东栎次生林各植物层（乔木层、林下植物层（草本和灌丛）、枯落物层）生物量随郁闭度及林分密度的变化,确定能使各层生物量均维持在较高水平的最佳林分结构,以充分发挥森林多种功能。 方法 基于野外调查样地数据,利用上外包线分析法确定辽东栎林各层生物量对林分密度和郁闭度的响应规律,并经综合权衡后确定最佳郁闭度范围。 结果 随郁闭度和林分密度增加,乔木层生物量先快速增加,在郁闭度0.75和密度1 050株∙hm−2时分别达到292.3和296.9 t∙hm−2,分别为其外包线最大值的80%和82%,之后增速变缓;林下植物层生物量先缓慢下降,在郁闭度0.75和密度1 150株∙hm−2时分别达到9.08和9.17 t∙hm−2,分别为其外包线最大值的89%和86%,之后快速减小;枯落物层生物量先逐渐增加,在郁闭度0.75时达到33.34 t∙hm−2,为外包线最大值的97%。综合分析各层生物量随郁闭度的变化表明,需维持林冠郁闭度在0.75～0.79左右,才能使林分各层生物量都保持在各自上外包线最大值的80%以上。 结论 在宁夏六盘山区,要使辽东栎林的乔木层、林下植物层、枯落物层的生物量均维持在较高水平,需维持林冠郁闭度在最佳范围0.75～0.79左右,这可用于指导该地区及类似地区的辽东栎林多功能经营。     

Wood net primary production resilience in an unmanaged forest transitioning from early to middle succession

The mixed deciduous forests of the upper Midwest, USA are approaching an ecological threshold in which early successional canopy trees are reaching maturity and beginning to senesce, giving way to a more diverse canopy of middle and late successional species. The net primary production (NPP) of these forests is generally considered past peak and in decline, but recent studies show a striking resilience in the NPP trajectories of some middle and late successional forests; yet, the mechanisms controlling such temporal changes in NPP are largely unknown. At the University of Michigan Biological Station in northern Michigan, we used a ≥9-year continuous record of wood net primary production (NPP), leaf area index (LAI), canopy composition, and stem mortality in 30 forested plots to identify the constraints on wood NPP as a mixed forest transitions from early to middle succession. Although wood NPP decreased over time in most stands, the rate of decline was attenuated when the canopy comprised a more diverse assemblage of early and middle/late successional species. The mechanism for sustained NPP in stands with more species diverse canopies was the proliferation of LAI by intact later successional tree species, even as stem mortality rates of early successional trees increased. We conclude that projections of carbon sequestration for the aging mixed forests of the upper Midwest should account for species composition shifts that affect the resilience wood NPP.     

Understory vegetation response after 30 years of interval prescribed burning in two ponderosa pine sites in northern Arizona,USA

Southwestern USA ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) forests evolved with frequent surface fires and have changed dramatically over the last century. Overstory tree density has sharply increased while abundance of understory vegetation has declined primarily due to the near cessation of fires. We examined effects of varying prescribed fire-return intervals (1, 2, 4, 6, 8, and 10 years, plus unburned) on the abundance and composition of understory vegetation in 2007 and 2008 after 30+ years of fall prescribed burning at two ponderosa pine sites. We found that after 30 years, overstory canopy cover remained high, while understory plant canopy cover was low, averaging <12% on all burn intervals. We attributed the weak understory response to a few factors – the most important of which was the high overstory cover at both sites. Graminoid cover and cover of the major grass species, Elymus elymoides (squirreltail), increased on shorter fire-return intervals compared to unburned plots, but only at one site. Community composition differed significantly between shorter fire-return intervals and unburned plots at one site, but not the other. For several response variables, precipitation levels appeared to have a stronger effect than treatments. Our findings suggest that low-severity burn treatments in southwestern ponderosa pine forests, especially those that do not decrease overstory cover, are minimally effective in increasing understory plant cover. Thinning of these dense forests along with prescribed burning is necessary to increase cover of understory vegetation.     

Response of aspen stands to forest tent caterpillar defoliation and subsequent overstory mortality in northeastern Ontario,Canada

Overstory mortality, understory tree recruitment, and vegetation development were assessed in trembling aspen (Populus tremuloides Michx.) stands following two recent episodes of forest tent caterpillar defoliation (Malacosoma disstria Hbn.) in northeastern Ontario. The results suggest that poplar (aspen and balsam poplar (Populus balsamifera L.)) mortality increased with consecutive years of insect defoliation occurring from the mid-1980s to mid-2000s and the proportion of poplars in the overstory, but decreased with improved pre-defoliation tree vigour (DBH increment). The first outbreak, which lasted from the mid-1980s to early 1990s, was more severe in terms of insect defoliation and contributed more to poplar mortality and decline. The decline began in the late 1990s and peaked in early 2000s. Poplar regeneration and understory shrubs responded rapidly to foliage loss to insect defoliation and mortality of overstory poplars. The regenerated poplars were able to maintain their growth under developing shrubs and residual overstory canopy and numbers were sufficient to compensate for the poplar trees lost to insect infestation. The defoliation-induced overstory decline will accelerate the transition of aspen stands to conifer dominance through enhanced conifer recruitment and growth, and reduced hardwood overstory in aspen-dominated stands, while hardwood dominance will persist in pure aspen stands. From a timber supply perspective, the decline caused by forest tent caterpillar defoliation could delay the availability of aspen stands for harvesting by 40–50 years.     

Stand structure, fuel loads, and fire behavior in riparian and upland forests, Sierra Nevada Mountains, USA; a comparison of current and reconstructed conditions

Fire plays an important role in shaping many Sierran coniferous forests, but longer fire return intervals and reductions in area burned have altered forest conditions. Productive, mesic riparian forests can accumulate high stem densities and fuel loads, making them susceptible to high-severity fire. Fuels treatments applied to upland forests, however, are often excluded from riparian areas due to concerns about degrading streamside and aquatic habitat and water quality. Objectives of this study were to compare stand structure, fuel loads, and potential fire behavior between adjacent riparian and upland forests under current and reconstructed active-fire regime conditions. Current fuel loads, tree diameters, heights, and height to live crown were measured in 36 paired riparian and upland plots. Historic estimates of these metrics were reconstructed using equations derived from fuel accumulation rates, current tree data, and increment cores. Fire behavior variables were modeled using Forest Vegetation Simulator Fire/Fuels Extension.Riparian forests were significantly more fire prone under current than reconstructed conditions, with greater basal area (BA) (means are 87 vs. 29 m²/ha), stand density (635 vs. 208 stems/ha), snag volume (37 vs. 2 m³/ha), duff loads (69 vs. 3 Mg/ha), total fuel loads (93 vs. 28 Mg/ha), canopy bulk density (CBD) (0.12 vs. 0.04 kg/m³), surface flame length (0.6 vs. 0.4 m), crown flame length (0.9 vs. 0.4 m), probability of torching (0.45 vs. 0.03), predicted mortality (31% vs. 17% BA), and lower torching (20 vs. 176 km/h) and crowning indices (28 vs. 62 km/h). Upland forests were also significantly more fire prone under current than reconstructed conditions, yet changes in fuels and potential fire behavior were not as large. Under current conditions, riparian forests were significantly more fire prone than upland forests, with greater stand density (635 vs. 401 stems/ha), probability of torching (0.45 vs. 0.22), predicted mortality (31% vs. 16% BA), and lower quadratic mean diameter (46 vs. 55 cm), canopy base height (6.7 vs. 9.4 m), and frequency of fire tolerant species (13% vs. 36% BA). Reconstructed riparian and upland forests were not significantly different. Our reconstruction results suggest that historic fuels and forest structure may not have differed significantly between many riparian and upland forests, consistent with earlier research suggesting similar historic fire return intervals. Under current conditions, however, modeled severity is much greater in riparian forests, suggesting forest habitat and ecosystem function may be more severely impacted by wildfire than in upland forests.     

Morphological acclimation and growth of ash (Fraxinus pennsylvanica Marsh.) advance regeneration following overstory harvesting in a Mississippi River floodplain forest

Stand-level growth responses and plant-level patterns of biomass accumulation and distribution were examined to learn how stand structure influences morphological acclimation and growth of green ash (Fraxinus pennsylvanica Marsh.) advance regeneration following overstory harvesting. Nine, 20-ha plots that received clearcut harvesting (100% basal area removal), partial harvesting (50% basal area removal), or no harvesting (control) were sampled to measure height, root-collar diameter, leaf, stem and root biomass, and leaf mass ratio (LMR), stem mass ratio (SMR) and root mass ratio (RMR) of ash regeneration. Six years after treatment, plot-level analyses indicated that ash growth was greatest in plots receiving clearcut harvesting, and least in control plots. Examination of LMR, SMR and RMR revealed that this growth response was not associated with acclimation that altered plant morphology. Total biomass ranged 275-fold among sampled plants, and much of this variation was accounted for by measurements of stand leaf area index (LAI). Along the gradient of stand LAI, values greater than 2 inhibited biomass accumulation. Stand LAI values less than 1.5 promoted ash biomass accumulation which reached a maximum where LAI values approached 0.7 and tapered above or below this value. Our findings indicate that green ash regeneration can be managed beneath light canopy cover, and the ability of seedlings to establish and persist beneath closed canopies and vigorously respond to release without having to endure prolonged morphological acclimation provides flexibility in developing regeneration protocols.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.