Conifer expansion reduces the competitive ability and herbivore defense of aspen by modifying light environment and soil chemistry

Disturbance patterns strongly influence plant community structure. What remains less clear, particularly at a mechanistic level, is how changes in disturbance cycles alter successional outcomes in plant communities. There is evidence that fire suppression is resulting in longer fire return intervals in subalpine forests and that these lengthened intervals increase competitive interactions between aspen and conifer species. We conducted a field and greenhouse study to compare photosynthesis, growth and defense responses of quaking aspen and subalpine fir regeneration under light reductions and shifts in soil chemistry that occur as conifers increase in dominance. The studies demonstrated that aspen regeneration was substantially more sensitive to light and soil resource limitations than that of subalpine fir. For aspen, light reductions and/or shifts in soil chemistry limited height growth, biomass gain, photosynthesis and the production of defense compounds (phenolic glycosides and condensed tannins). Biomass gain and phenolic glycoside concentrations were co-limited by light reduction and changes in soil chemistry. In contrast, subalpine fir seedlings tended to be more tolerant of low light conditions and showed no sensitivity to changes in soil chemistry. Unlike aspen, subalpine fir increased its root to shoot ratio on conifer soils, which may partially explain its maintenance of growth and defense. The results suggest that increasing dominance of conifers in subalpine forests alters light conditions and soil chemistry in a way that places greater physiological and growth constraints on aspen than subalpine fir, with a likely outcome being more successful recruitment of conifers and losses in aspen cover.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Drought and fire suppression lead to rapid forest composition change in a forest-prairie ecotone

Altered fire regimes and increased drought can lead to major vegetation changes, especially in ecotones. A decrease in fire can lead to woody species encroachment in prairies and increasing forest stand density. The threat of global climate change raises questions about potential increases in the length, severity, and incidence of droughts substantially altering species composition. Re-measured upland forests in south-central North America's midcontinent forest-prairie ecotone exhibited major changes in woody species composition and structure over fifty years and successional trajectories appeared to favor invasive Juniperus virginiana L. over the previous dominant Quercus species. The objective of this study was to determine whether climate and fire exclusion affected the recruitment history of dominant woody species in these upland forests located near the xeric western edge of the eastern deciduous forest biome of North America. We removed cores and cross-sections from 992 J. virginiana, Quercus marilandica Münchh. and Q. stellata Wangenh. trees from eleven forest stands located across central and northwest Oklahoma, and determined their ages using standard dendrochronological methods. Recruitment of all species increased following a severe mid-20th century drought, but a rapid increase in J. virginiana recruitment and decrease in Quercus recruitment appeared to be linked to a decrease in fire. Future fire regime changes and increased drought due to global climate change could lead to widespread shifts from Quercus- to Juniperus- dominated forests and cause substantial changes to ecosystem services.     

Effect of fire severity and site slope on diversity and structure of the ectomycorrhizal fungal community associated with post-fire regenerated Pinus pinaster Ait. seedlings

We investigated the diversity and structure of the ectomycorrhizal (EM) fungal community associated with post-fire regenerated Pinus pinaster Ait., and the influence of fire severity and site slope on EM assemblage patterns. Seedlings were sampled in the first autumn and in both spring and autumn of the second growing season after fire, in a total of three samplings. EM percentages per seedling were assessed, morphotypes described, and tentative identification of EM types performed by restriction fragment length polymorphism (RFLP) and sequencing of nrDNA internal transcribed spacer (ITS) region. Seedlings were highly mycorrhizal in all samplings, independently of the factors studied. A total of 45 EM types were identified, and richness and diversity significantly increased from the first to the second autumn after fire. Neither fire severity nor slope had a significant effect on fungal richness and diversity. Overall EM community composition was similar in all samplings, although fire severity, site slope and elapsed time after fire caused evident shifts in presence or in relative frequencies of a number of EM types. No significant effect of fire severity or slope on EM assemblage patterns was detected in the first two samplings after fire. However, a significant effect of fire severity was observed at the end of the second growing season. The harvest of burned wood did not significantly affect EM fungal assemblages although the slope did. We conclude that there was a high potential of active EM inoculum in soil immediately after fire colonizing post-fire natural regenerated P. pinaster seedlings with high EM percentages, and that factors defining burn intensity, such as fire severity and topography, directly influenced the species composition and assemblage patterns of EM fungal communities, triggering replacements and succession of EM fungal species.     

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.     

Characteristics of aspen infected with heartrot: Implications for cavity-nesting birds

Phellinus tremulae is an important fungal decay agent common to aspen and a critical component to the cavity-nesting bird complex found in western aspen stands. Little information exists on the conditions that facilitate infection and spread of P. tremulae in aspen forests. I used Forest Inventory and Analysis (FIA) data to explore the relationships of several tree and stand characteristics to the presence and frequency of P. tremulae in aspen measured across several western states of the United States. Results suggest a strong relationship between tree age, tree diameter, and compacted crown ratio with infection frequency in trees while stand purity, canopy cover and stand age had a positive relationship with the occurrence of P. tremulae in forest stands containing aspen. Logistic regression modeling identified stand age as the only variable that increased the odds of predicting infection at the stand-level while all tree-level variables were included in the tree model. Data also show that infection rates in the study area were lower than in other parts of aspen's range, and that average size of infected trees was smaller in the study area than those reported elsewhere. These results have important implications to management of aspen for wildlife, especially for birds that use decayed aspen for nesting.     

Ectomycorrhizal communities associated with silver fir seedlings (Abies alba Mill.) differ largely in mature silver fir stands and in Scots pine forecrops

Context

The requirement for rebuilding forecrop stands besides replacement of meadow vegetation with forest plants and formation of soil humus is the presence of a compatible ectomycorrhizal (ECM) fungal community.

Aims

This study aims to assess ectomycorrhizal fungi diversity associated with silver fir (Abies alba Mill.) seedlings regenerating in silver fir stands and Scots pine forecrops.

Methods

One-year-old seedlings were sampled in six study sites: three mature fir forests and three pine forests. ECM fungi were identified by polymerase chain reaction amplification and sequencing of the internal transcribed spacer of rDNA.

Results

The mean mycorrhizal colonization exceeded 90 %. Thirty-six ectomycorrhizal taxa were identified in fir stands and 23 in pine forecrops; ten out of these species were common to both stands. The fungal communities were different between study sites (R?=?0.1721, p?=?0.0001). Tomentella stuposa was the only species present at all sites.

Conclusion

Silver fir seedlings in Scots pine forecrops supported smaller ECM fungal communities than communities identified in mature silver fir stands. Nevertheless, fungal colonization of seedling roots was similar in both cases. This suggests that pine stands afforested on formerly arable land bear enough ECM species to allow survival and growth of silver fir seedlings.     

Quantifying successional rates in western aspen woodlands: Current conditions,future predictions

Stands of quaking aspen (Populus tremuloides) rank among the most biologically diverse plant communities across the intermountain region of western North America. Marked declines of aspen have occurred in recent decades, likely due to a combination of effects from changes in fire regimes, herbivory, climate (e.g. drought), and interspecific competition with conifer species. However, it is poorly understood how the effects of these factors are manifested at a landscape scale over decadal time periods. Analysis of field data combined with topographic information collected across the 500,000 ha Owyhee Plateau in southwestern Idaho revealed that aspen in the area occur in three different biophysical settings; First, aspen stands exist at high altitudes on south-facing slopes where local conifer species are not likely to occur because of limiting temperature or precipitation levels under current climate conditions. In these areas aspen is the potential vegetation type rather than conifers. Second, aspen grow on anomalously wet microsites (e.g. near springs), and third, aspen grow within upland mixed aspen/conifer stands, which are experiencing rapid rates of conifer establishment. Based on a paired t-test (α = 0.05) we conclude that stands growing on wet microsites show significantly slower successional rates of conifer establishment relative to upland aspen stands. We developed a conceptual state-and-transition model for upland aspen/conifer stands occurring across a range of topographic positions. We then parameterized the model using extensive field data in the vegetation dynamics computer simulation model Vegetation Dynamics Development Tool (VDDT), and examined the current and future aspen distribution under varying fire regimes. Model results indicate that average fire return intervals of 50–70 years are desirable for maintenance of aspen in upland areas where conifers are present. Under the current fire regime in the area many upland aspen/conifer stands will likely be lost within 80–200 years. Thresholds for the effect of conifer encroachment and browsing on aspen regeneration identified through this research are similar to those described by others across the West. We therefore suggest that the results presented for the Owyhee Plateau are likely applicable to semi-arid aspen woodlands across the American West where succession to conifers is a cause of aspen decline.     

Historical fire and vegetation dynamics in dry forests of the interior Pacific Northwest,USA, and relationships to Northern Spotted Owl (Strix occidentalis caurina) habitat conservation

Regional conservation planning frequently relies on general assumptions about historical disturbance regimes to inform decisions about landscape restoration, reserve allocations, and landscape management. Spatially explicit simulations of landscape dynamics provide quantitative estimates of landscape structure and allow for the testing of alternative scenarios. We used a landscape fire succession model to estimate the historical range of variability of vegetation and fire in a dry forest landscape (size ca. 7900 km²) where the present-day risk of high severity fire threatens the persistence of older closed canopy forest which may serve as Northern Spotted Owl (Strix occidentalis caurina) habitat. Our results indicated that historically, older forest may have comprised the largest percentage of the landscape (∼35%), followed by early successional forest (∼25%), with about 9% of the landscape in a closed canopy older forest condition. The amount and condition of older forest varied by potential vegetation type and land use allocation type. Vegetation successional stages had fine-grained spatial heterogeneity in patch characteristics, with older forest tending to have the largest patch sizes among the successional stages. Increasing fire severities posed a greater risk to Northern Spotted Owl habitat than increasing fire sizes or frequencies under historical fire regimes. Improved understanding of historical landscape-specific fire and vegetation conditions and their variability can assist forest managers to promote landscape resilience and increases of older forest, in dry forests with restricted amounts of habitat for sensitive species.     

Water use by whitebark pine and subalpine fir: potential consequences of fire exclusion in the northern Rocky Mountains

In subalpine forests of the northern Rocky Mountains, fire exclusion has contributed to large-scale shifts from early-successional whitebark pine (Pinus albicaulis Engelm.) to late-successional subalpine fir (Abies lasiocarpa (Hook.) Nutt.), a species assumed to be more shade tolerant than whitebark pine and with leaf to sapwood area ratios (A(L):A(S)) over twice as high. Potential consequences of high A(L):A(S) for subalpine fir include reduced light availability and, if hydraulic sufficiency is maintained, increased whole-tree water use. We measured instantaneous gas exchange, carbon isotope ratios and sap flow of whitebark pine and subalpine fir trees of different sizes in the Sapphire Mountains of western Montana to determine: (1) whether species-specific differences in gas exchange are related to their assumed relative shade tolerance and (2) how differences in A(L):A(S) affect leaf- and whole-tree water use. Whitebark pine exhibited higher photosynthetic rates (A = 10.9 micromol x m(-2) x s(-1) +/- 1.1 SE), transpiration rates (E = 3.8 mmol x m(-2) x s(-1) +/- 0.7 SE), stomatal conductance (g(s) = 166.4 mmol x m(-2) x s(-1) +/- 5.3 SE) and carbon isotope ratios (delta13C = -25.5 per thousand +/- 0.2 SE) than subalpine fir (A = 5.7 micromol x m(-2) x s(-1) +/- 0.9 SE; E = 1.4 mmol x m(-2) x s(-1) +/- 0.3 SE; g(s) = 63.4 mmol x m(-2) x s(-1) +/- 1.2 SE, delta13C = -26.2 per thousand +/- 0.2 SE; P < 0.01 in all cases). Because subalpine fir had lower leaf-area-based sap flow than whitebark pine (QL = 0.33 kgx m(-2) x day(-1) +/- 0.03 SE and 0.76 kg x m(-2) x day(-1) +/- 0.06 SE, respectively; P < 0.001), the higher A(L):A(S) in subalpine fir did not result in direct proportional increases in whole-tree water use, although large subalpine firs used more water than large whitebark pines. The linear relationships between tree size and daily water use (r2 = 0.94 and 0.97 for whitebark pine and subalpine fir, respectively) developed at the Sapphire Mountains site were applied to trees of known size classes measured in 12 natural subalpine stands in the Bob Marshall Wilderness Complex (western Montana) ranging from 67 to 458 years old. Results indicated that the potential for subalpine forests to lose water by transpiration increases as succession proceeds and subalpine fir recruits into whitebark pine stands.     

Successional process of Picea crassifolia forest after logging disturbance in semiarid mountains: A case study in the Qilian Mountains,northwestern China

Selective logging is the most widely employed method of commercial timber production in Asia, and its impact on forest structure, composition, and regeneration dynamics is considerable. However, the successional processes in forest communities after logging in semiarid mountains are poorly understood. To provide more information on these processes, we used data from tree rings, direct and indirect age determinations, and field measurements of stand structure to reconstruct the historical disturbance regime, stand development patterns, and successional processes in a natural Picea crassifolia forest community in the Qilian Mountains of northwestern China. The results showed that the density of P. crassifolia forest increased significantly after logging. The densities of second growth forests 30 and 70 years after logging disturbance had increased to 2874% and 294% of primary forest's density, respectively. Logging disturbance did not alter tree species composition of logged stands. However, the diversity of understory species changed significantly among the successional phases. Logging disturbance decreased the spatial heterogeneity of second growth forest. The spatial distributions of recruitment were affected by the location of the remaining trees. There was less recruitment near the remaining trees than near forest that had been cut. In addition, logging disturbance also induced a growth release for the trees on the sites sampled. Our results imply that the succession and regeneration of P. crassifolia forest may be improved if the remaining trees could be retained relative uniform distribution pattern, thinning or selective logging could be performed to height density, exotic shrubs could be removed or the shrubs cover could be reduced during the earlier successional stages.     

Using a retrospective dynamic competition index to reconstruct forest succession

Understanding forest dynamics and stand structures is crucial for predicting forest succession. However, many forests have been altered due to century-long land-use practices, which complicates the reconstruction of past and current successional trajectories. For a better understanding of successional processes, we suggest studying the intra- and interspecific competition among single trees across time. We introduce a tree-ring based competition index to reconstruct the competitive dynamics of individual trees over time. This new retrospective dynamic competition index combines a temporal and a spatial component by calculating the yearly ratio between the basal area increments (bai) of the neighbouring trees and the subject tree. The new index is applied to mixed Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) stands in the inner-Alpine dry-valley Valais, for which a change in species composition is hypothesised. The aim is to analyse current stand structures in terms of recent changes in the competitive interactions at the single tree level and to relate these competitive dynamics to land-use change and increasing drought due to climate change. On five plots, the positions of 456 trees were recorded and increment cores were taken to derive bai data. The individual dynamic competition index curves were aggregated in clusters, which define typical patterns of competitive dynamics in both tree species. A large percentage of the trees (87% in oak, 70% in pine) were clustered into a group of trees with constant competition at a relatively low level. However, a smaller group of pines (20%) had recently faced increasing competition. In addition, stand structure analyses indicated a change towards a higher proportion of oak. This change in the competitive ability between oak and pine was found to be related to drought, in that oak had a competitive advantage in dry years. Furthermore, the high proportion of dead branches in pines with decreasing competitive abilities indicated increasing competition for light as a consequence of natural development towards a later successional stage that favours the more shade-tolerant oak. The new retrospective dynamic competition index proved to be promising in studying forest succession. The tree-ring based method allows us to identify changes in the competitive ability of single trees with a high temporal resolution and without repeated assessments.     

Above- and below-ground biomass and nutrient distribution of a paper birch and subalpine fir mixed-species stand in the Sub-Boreal Spruce zone of British Columbia

Stand structure, height and diameter growth, above- and below-ground biomass, and nutrient concentrations and content were determined for a 35-year-old fire-origin paper birch (Betula papyrifera) and subalpine fir (Abies lasiocarpa) mixed-species stand in the Sub-Boreal Spruce (SBS) zone of British Columbia. Paper birch, which formed the dominant overstory following the 1961 fire, had normal distributions of height and diameter classes. Subalpine fir, which dominated the understory, had the reverse J-shaped height and diameter distributions that are expected of a shade tolerant, climax species. Paper birch grew more than three times the height of subalpine fir. Growing in the summer shade of the birch, subalpine fir had slow but steady height growth during the first 10–15 years, after which height growth declined somewhat. Allometric equations, relating dry weight of foliage, branches, stemwood, stembark, roots, and total biomass to diameter at breast height (DBH), were developed to estimate above- and below-ground biomass. Total biomass of paper birch reached 83.2 t ha⁻¹, while subalpine fir biomass was 26.7 t ha⁻¹. Subalpine fir allocated more biomass to foliage and branches compared to paper birch. Foliage of paper birch had higher nutrient concentrations of N, P, K, Ca, and Mg than subalpine fir foliage. Branches and stembark of subalpine fir had higher P, and Ca concentrations than paper birch. Subalpine fir branches contained more of all examined nutrients than paper birch branches. This is a significant component in nutrient cycling of the mixed-species forest.     

Stand structure and successional trends in virgin boreal forest reserves in Sweden

Fire history and stand structure was examined in twelve virgin forest stands situated within forest reserves in northern Sweden. The selected stands represented fire refuges as well as different successional stages after fire. Six of the stands were dominated by Norway spruce (Picea abies L. Karst.), three were dominated by Scots pine (Pinus sylvestris L.), and three were dominated by hairy birch (Betula pubescens Ehrh.) or aspen (Populus tremula L.). In 3 of the southernmost stands, the average fire interval was 34 to 65 years during the late 1600s to late 1800s, but since 1888 no fires had occurred in any of the stands. The absence of fire disturbance since 1888 is probably caused by the fire suppression in the overall landscape. The standing volume of living trees ranged between 87 and 511 m³ ha⁻¹ while the volume of dead trees, including both snags and logs, ranged between 27 and 201 m³ ha⁻¹. The volume of dead trees constituted ca. 30% of the total stem volume. In the conifer dominated stands, there was a statistically significant relationship between total stem volume, including both living and dead trees, and site productivity. A comparison between the amount of dead and living trees indicated substantial changes in tree species composition in several stands. It is suggested that data on the amount of dead trees, especially logs, and its distribution over decay classes could be used to examine the continuity of certain tree species. All stands had a multi-sized tree diameter distribution, which in most cases was similar to a reversed J-shaped distribution. In general spruce was numerous in the seedling cohort and in small diameter classes, indicating that its proportion in the stands was stable, or was increasing at the expense of pioneer tree species such as pine, aspen and silver birch (Betula pendula Roth.). The most numerous species in the seedling cohort, rowan (Sorbus aucuparia L.), was almost totally missing in the tree layer, indicating a high browsing pressure preventing rowan seedlings from growing into trees. The general increase of spruce and the sparse regeneration of pioneer species, in the stands previously affected by fire, are discussed in relation to natural disturbance regimes, biological diversity and nature conservation policies. It is proposed that reintroduction of fire disturbance is a necessity for future management plans of forest reserves. Other management practices to increase species diversity within forest reserves are also discussed.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Soil CO2 efflux in an Afromontane forest of Ethiopia as driven by seasonality and tree species

Variability of soil CO₂ efflux strongly depends on soil temperature, soil moisture and plant phenology. Separating the effects of these factors is critical to understand the belowground carbon dynamics of forest ecosystem. In Ethiopia with its unreliable seasonal rainfall, variability of soil CO₂ efflux may be particularly associated with seasonal variation. In this study, soil respiration was measured in nine plots under the canopies of three indigenous trees (Croton macrostachys, Podocarpus falcatus and Prunus africana) growing in an Afromontane forest of south-eastern Ethiopia. Our objectives were to investigate seasonal and diurnal variation in soil CO₂ flux rate as a function of soil temperature and soil moisture, and to investigate the impact of tree species composition on soil respiration. Results showed that soil respiration displayed strong seasonal patterns, being lower during dry periods and higher during wet periods. The dependence of soil respiration on soil moisture under the three tree species explained about 50% of the seasonal variability. The relation followed a Gaussian function, and indicated a decrease in soil respiration at soil volumetric water contents exceeding a threshold of about 30%. Under more moist conditions soil respiration is tentatively limited by low oxygen supply. On a diurnal basis temperature dependency was observed, but not during dry periods when plant and soil microbial activities were restrained by moisture deficiency. Tree species influenced soil respiration, and there was a significant interaction effect of tree species and soil moisture on soil CO₂ efflux variability. During wet (and cloudy) period, when shade tolerant late successional P. falcatus is having a physiological advantage, soil respiration under this tree species exceeded that under the other two species. In contrast, soil CO₂ efflux rates under light demanding pioneer C. macrostachys appeared to be least sensitive to dry (but sunny) conditions. This is probably related to the relatively higher carbon assimilation rates and associated root respiration. We conclude that besides the anticipated changes in precipitation pattern in Ethiopia any anthropogenic disturbance fostering the pioneer species may alter the future ecosystem carbon balance by its impact on soil respiration.     

Composition and structure of natural mixed-oak stands in northern and central Portugal

Stand composition and structure of natural mixed-oak stands of common-oak (Quercus robur L.) and pyrenean-oak (Quercus pyrenaica Willd.) were studied. Diverse compositional and structural elements in early and late successional stand stages were analysed. The study was conducted in north and central Portugal where different natural mixed oak forests types are located. The following mixed-oak forest types involving common-oak and pyrenean-oak were studied: common-oak & other hardwoods; common-oak & cork-oak (Quercus suber L.); ash (Fraxinus angustifolia Vahl) & pyrenean-oak; and pyrenean-oak & madrone (Arbutus unedo L.). Measurements were made in early and late successional stand stages on the different mixed oak forest types. Different stand characteristics and indices were used to describe and compare stand structure and composition. The study showed changes in species diversity and stand structure. Most tree species in mature stands are present in early stages but with higher abundance. Shannon diversity index may change between 0.798 and 1.915. Significant differences on species diversity and abundance were found depending on the forest type and successional stage. Mature mixed-oak forests have high species diversity with an abundance of small to medium tree size species. Species distribution and diameter differentiation indices range from 0.30 to 0.70 and 0.52 to 0.82, respectively, revealing significant structural complexity. The average number of standing and downed dead trees was 265 and 83 trees ha⁻¹ for early and late stage, respectively, with 6.9 and 65.4 m³ ha⁻¹. Higher values of stand diversity index were 41 and 53 in more complex and developed forests. Later stand stages have complex structure, with a wider range of tree diameter distribution and higher degree of irregularity.     

Diversity and succession of adventive and indigenous vascular understorey plants in Pinus radiata plantation forests in New Zealand

The vegetation of Pinus radiata plantation forests in New Zealand was studied to examine how the indigenous flora has responded to this novel habitat. A chronosequence of stands about 5, 16 and 27 years was assessed in each of four different biogeographic regions to test the effects of several stand and site factors on the succession of vascular understorey plant communities. A total of 202 indigenous and 70 adventive vascular plant species were found across all study areas, with considerable geographic variation among forests in species composition, species richness (range 48–135 species), and the percentage of indigenous species (50–86%). Both richness and cover of adventive species decreased significantly over time, whereas richness and cover of indigenous species was highest in the oldest stands, and overall species richness was lowest at mid-rotation. The guild composition changed from dominance of grasses and forbs in young stands to dominance of ferns and understorey trees in mature stands. These temporal changes were accompanied by a decrease in light-demanding pioneer species and an increase in shade tolerant, later seral species adapted to a forest environment. Measurements of the degree of canopy closure in stands with low or high stocking and modelling of temporal changes of canopy closure indicated that these understorey plant dynamics are influenced by changes in light availability as stands age. Despite the successional changes within forests, geographic variation more strongly influenced understorey communities because stands within a forest area were grouped together in DCA and TWINSPAN analyses, along rainfall and temperature gradients. Although the canopy species of such intensively managed plantation forests is an alien element in the New Zealand flora, the sheltered forest environment of older stands allows the establishment of a mostly indigenous forest understorey community with considerable similarities to indigenous forests located nearby.     

Do position and species identity of neighbours matter in 8–15-year-old post harvest mesic stands in the boreal mixedwood?

Neighbourhood competition indices (NCI), where position and species identity of neighbours are known, have been used to investigate growth and competitive interactions among adult trees. In this study, we used NCI in 8–15-year-old stands following clear-cutting in a boreal mixedwood forest of eastern Canada to improve our understanding of early successional forest dynamics. Trees of increasing diameter from the center (≥1 cm) to the edge (≥5 cm) were mapped in twenty-five circular 450 m² plots. Target trees (DBH ≥ 1 cm) were sampled in plot center to determine their annual radial stem growth. For each species, we compared a set of growth models using either a spatially explicit NCI or a non-spatial competition index. Both types of indices estimated a species-specific competition coefficient for each pair of competitor–target species. NCI were selected as the best competition model for all target species although differences in variance explained relative to the non-spatial index were small. This likely indicates that competition occurs at the local level but that the high density and the relative uniformity of these young stands creates similar neighbourhoods for most trees in a given stand. The effective neighbourhood radius for competitors varied among species and was smaller for shade tolerant species. Intraspecific neighbours were the strongest competitors for most species. Aspen (Populus tremuloides) was a weak competitor for all species as opposed to balsam fir (Abies balsamea) which was a strong competitor in all cases. These results are in contradiction with some widely used forest policies in North America (e.g. free-to-grow standards) that consider broadleaf species, such as aspen, as the strongest competitors. For these early successional forests, the decision regarding the use of spatial or non-spatial competition indices should rest on the intended use. For even-age management, spatial indices might not justify their use in high-density stands but they are needed for the simulation of novel harvest techniques creating complex stand structure.