Radial and stand‐level thinning treatments: 15‐year growth response of legacy ponderosa and Jeffrey pine trees

Restoration efforts to improve vigor of large, old trees and decrease risk to high‐intensity wildland fire and drought‐mediated insect mortality often include reductions in stand density. We examined 15‐year growth response of old ponderosa pine (Pinus ponderosa) and Jeffrey pine (Pinus jeffreyi) trees in northeastern California, U.S.A. to two levels of thinning treatments compared to an untreated (control) area. Density reductions involved radial thinning (thinning 9.1 m around individual trees) and stand thinning. Annual tree growth in the stand thinning increased immediately following treatment and was sustained over the 15 years. In contrast, radial thinning did not increase growth, but slowed decline compared to control trees. Available soil moisture was higher in the stand thinning than the control for 5 years post‐treatment and likely extended seasonal tree growth. Our results show that large, old trees can respond to restoration thinning treatments, but that the level of thinning impacts this response. Stand thinning must be sufficiently intensive to improve old tree growth and health, in part due to increasing available soil moisture. Importantly, focusing stand density reductions around the immediate neighborhood of legacy trees was insufficient to elicit a growth response, calling into question treatments attempting to increase vigor of legacy trees while still maintaining closed canopies in dry, coniferous forest types. Although radial thinning did not affect tree growth rates, this treatment may still achieve other resource objectives not studied here, such as protecting wildlife habitat, reducing the risk of severe fire injury, and decreasing susceptibility to bark beetle attacks.     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

From Pine Plantations to Natural Stands. Ecological Restoration of a <Emphasis Type="Italic">Pinus canariensis</Emphasis> Sweet,ex Spreng forest

We evaluated silvicultural thinning of pine plantations in order to determine the extent to which plantations treated in this way showed a greater structural similarity to natural stands. Specifically, we tested for differences in community structure (increase of DBH, increase of height and canopy height) and regeneration (seedlings and saplings <1, 1–2 and >2 years old) in response to thinning treatments (20% and 50% removal of density). We compared the variables of the thinned plots with those of the control plots (no thinning of living trees). Comparison of the structural variables between any treatments is of limited value due to the high intra- and inter-plot environmental variability (both slope and orientation affect tree growth to a significant degree). We therefore used ordination methods (Redundancy Analysis, RDA) to monitor covariation and to select non-redundant explanatory variables. We tested for differences between control and managed plots using Monte Carlo tests for the eigenvalues of the obtained axis of the RDA. Of the two treatments, only the 50% thinning treatment was significantly different from the control plots (in which only dead pines were thinned). In ten years, the basal area of pines showed a 10% increase in 50% thinned plots in comparison with the control plots. The number of saplings >2 years old was also significantly higher in 50% thinned plots. The control plots typically had an appreciably higher density of dead trees and a greater number of seedlings. Fifty percent thinning is having a positive effect on the naturalization of the stand but subsequent management will be needed to ensure establishment of advance regeneration.     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Mechanical Restoration of California Mixed‐Conifer Forests: Does it Matter Which Trees Are Cut?

The montane ecosystems of northern California have been subjected to repeated manipulation and active fire suppression for over a century, resulting in changes in community structure that contribute to increased wildfire hazard. Ecosystem restoration via reduction of stand density for wildfire hazard mitigation has received substantial attention in recent years; however, many ecological questions remain unanswered. This study compares belowground effects of two alternative forest thinning treatments designed to restore the large, old tree component of late‐seral structure, one of which focuses on restoring Pinus ponderosa dominance (Pine‐preference) and the other of which promotes development of large trees regardless of species (Size‐preference). We evaluated forest floor and soil chemical and microbial parameters in six experimental thinning treatment units of 40 ha each in the Klamath National Forest of northern California 5–6 years after thinning. Inorganic N availability, soil organic C content, phenol oxidase activity, and forest floor C:N ratio were greater in the Size‐preference treatment, whereas forest floor N and soil pH were greater in the Pine‐preference treatment. Our results indicate that these two thinning strategies produce differences in the soil environment that has the potential to affect growth rates of trees that remain, as well as the growth and survivorship of newly established seedlings. Thus, which species/individuals are removed during structural restoration of these mixed‐conifer forests matters both to the belowground components of the ecosystem today and the vegetation and productivity of the ecosystem in future decades.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

外源性氮磷添加及林分密度对大叶相思林土壤酶活性的影响

研究了林分密度和外源性氮(N)磷(P)添加及二者的交互作用对土壤酶活性的影响,可以为当前不同密度的亚热带林在氮沉降和施磷肥条件下,影响我国亚热带人工林土壤养分转化的酶学机制提供理论依据。以大叶相思(Acacia auriculiformis)人工林为试验材料,选择氯化铵(NH_4Cl)和二水合磷酸二氢钠(NaH_2PO_4·2H_2O)作为N肥和P肥,氮磷添加设置4个处理,即对照、施N、施P和施N+P;种植密度设置低密度(1667株/hm~2)、中密度(2500株/hm~2)、较高密度(4444株/hm~2)和高密度(10000株/hm~2)4个水平。在4种密度的大叶相思人工林下各设置一个样地(即4个样地),每个样地内5个样方,每个样方内设置4种处理,合计共16个处理,5个重复,80个小区,测定土壤脲酶、磷酸酶和过氧化氢酶活性,分析林分密度和NP添加对大叶相思林地土壤酶活性的影响。研究结果表明:N添加对各林分密度下的土壤脲酶和过氧化氢酶活性均有抑制作用,P添加显著提高了过氧化氢酶活性和高密度林分大叶相思林地的土壤脲酶活性,却降低了低密度林分的土壤脲酶活性;土壤磷酸酶活性在P和N+P添加两种处理下显著小于对照和施N处理;土壤脲酶、磷酸酶以及对照和P添加处理下的过氧化氢酶活性均随着林分密度的升高呈现上升趋势,而土壤过氧化氢酶活性在N和N+P添加后随着林分密度的减小而升高。林分密度和外源性NP添加的交互作用对三种土壤酶的活性均有显著影响。     

Short‐term response of snowshoe hares to western larch restoration and seasonal needle drop

Old‐growth western larch has been degraded throughout much of its historic range due to extensive timber harvest and fire suppression. We examined the effects of a restoration treatment of western larch on snowshoe hares, a denizen of the boreal forest serving as a focal animal species to indicate the health of the restored ecosystem. We implemented a restoration treatment using “doughnut thinning” to accelerate development of old‐growth attributes in larch stands and simultaneously examined the short‐term effects on snowshoe hare density, survival, and movement. Although typical forest management activities tend to have adverse effects on hares especially in the short term, we found that the restoration treatment did not affect hare density or survival in the short term. In addition, despite significant decreases in cover coinciding with the larch needle drop, we found evidence of year‐round immigration into larch stands by hares suggesting larch stands are suitable year‐round hare habitat. Taken together, our findings suggest that a larch restoration treatment designed to accelerate the development of old‐growth attributes can be implemented so as to have no measurable short‐term detrimental effects on hares.     

Vegetation Structure,Composition, and Species Richness Across a 56‐year Chronosequence of Dry Tropical Forest on Providencia Island,Colombia1

We compared vegetation structure and species richness across a 56‐yr chronosequence of six replicated age classes of dry tropical forest on the island of Providencia, Colombia, in the Southwest Caribbean. Stand age classes were determined using sequential, orthorectified panchromatic aerial photos acquired between 1944 and 1996 and Landsat 7 ETM + satellite imagery from 2000. Along the chronosequence we established 59 plots of 2 × 50 m (0.01 ha) to document changes in species richness, basal area, tree height, stem density, and sprouting ability. All woody trees and shrubs >2.5 cm diameter at breast height (DBH) were censused and measured. Although woody species density reached a peak in stands from 32 to 56 yr old, rarefaction analysis showed that species richness increased linearly with stand age and was highest in stands 56 yr old or greater. Nonparametric, abundance‐based estimators of species richness also showed positive and linear associations with age. Basal area and mean tree height were positively associated with age since abandonment, while sprouting ability showed a negative relationship. Our results indicate rapid recovery of woody species richness and structural characteristics along this tropical dry forest chronosequence.     

Regeneration of a Coastal Pine (Pinus thunbergii Parl.) Forest 11 Years after Thinning,Niigata, Japan

To examine the effects of thinning intensity on wind vulnerability and regeneration in a coastal pine (Pinus thunbergii) forest, thinning with intensities of 20%, 30% and 50% was conducted in December 1997; there was an unthinned treatment as the control (total 8 stands). We re-measured the permanent sites to assess the regeneration characteristics 11 years after thinning. In the 50% thinned stand, seedlings aged from 2 to 10 years exhibited the highest pine seedling density and growth. The age composition ranged from 1–3 years with densities of 9.9 and 5.1 seedlings m⁻² in 30% and 20% thinned stands; only 1-year-old seedlings with a density of 6.1 seedlings m⁻² in the unthinned stand. Similar trends were found for the regeneration of broadleaved species such as Robinia pseudoacacia and Prunus serrulata. We speculate that the canopy openness and moss coverage contributed to the regeneration success in the 50% thinned stand, while the higher litter depth and lack of soil moisture induced the regeneration failure in the unthinned stand. The stands thinned at 20% or 30% were less favourable for pine regeneration than the stands thinned at 50%. Therefore, thinning with less than 30% canopy openness (20% and 30% thinned stands) should be avoided, and thinning at higher than 30% canopy openness (50% thinned stand, approximately 1500 stems ha⁻¹ at ages 40–50 years) is suggested for increasing regeneration in the coastal pine forest. The implications of thinning-based silviculture in the coastal pine forest management are also discussed. The ongoing development of the broadleaved seedlings calls for further observations.     

Succession of bird communities in young temperate rainforests following thinning

We repeated bird and vegetation surveys in 1991–1992 and 2005–2006 among young managed stands and old-growth forests in southeast Alaska to evaluate whether pre-commercial thinning of managed stands influenced the bird community. We compared decadal changes in bird densities and forest vegetation among 3 stand types: managed stands originating from clearcuts 35 years ago that were left untreated (unthinned), managed stands thinned at uniform spacing (thinned), and old growth with no prior timber harvest. We did not detect differences in decadal trends in avian densities between thinned and unthinned stands for 15 of 16 common bird species using a repeated-measures design. Thinning did not result in greater recruitment of overstory-nesting species as predicted. This was likely because of 1) similar increases in tree heights ( = 9–10 m) and canopy cover ( = 29–43%) between unthinned and thinned stands across decades and 2) the relatively young successional stage of these stands, which had only begun to recruit medium and large size conifers (dbh ≥ 36 cm). Decadal trends in densities of most (88%) understory-nesting bird species did not differ between thinned and unthinned stands. Shrub cover decreased by 22% and 31% across decades in thinned and unthinned stands, respectively. Bird community composition in managed stands reflected the general decadal changes in forest vegetation with a shift in dominance from understory species in the early 1990s (80–85% of total bird density) to an equal abundance of understory (45–54%) and overstory species in the mid-2000s. The latter was more similar to old-growth stands, which were dominated by overstory species (67–71%). Overstory-nesting birds in old growth increased in density by 49% across decades. Densities of cavity-nesting species remained unchanged in managed stands and less than densities in old growth across decades, possibly because of a lack of large trees and snags for nest sites. Overall, thinning of clearcut stands, the primary silvicultural system in the region, had few measurable benefits to birds nearly 20 years after treatment. Monitoring over the 70–100-year harvest rotation may be necessary to fully test whether thinning accelerates succession of bird communities in clearcut stands. However, partial harvests that retain large trees and snags should also be explored as alternatives to better maintain late-succession avifauna throughout the harvest rotation in southeast Alaska. © 2012 The Wildlife Society.     

Monitoring Landscape‐Scale Ponderosa Pine Restoration Treatment Implementation and Effectiveness

We evaluated landscape‐scale forest restoration treatment implementation and effectiveness in meeting objectives in a ponderosa pine forest at Mt. Trumbull, Arizona, U.S.A. The goal of the project was to alter forest structure by thinning and burning to more closely resemble forest conditions prior to Euro‐American settlement in 1870. We measured 117 permanent plots before (1996/1997) and after (2003) treatments. The plots were evenly distributed across the landscape (approximately 1,200 ha), about half of which was an untreated control. We evaluated treatment implementation and effectiveness based on 1870 structure and/or goals outlined by managers. The success of treatment implementation varied: about 94% of the area originally planned for restoration was treated in some manner by 2003, but only 70% received the full planned treatment (thin and burn). Although density of ponderosa pines >2.5 cm was reduced significantly by 66% from approximately 429 pines/ha to approximately 146 pines/ha in the treated area, the targeted residual density was exceeded by 111–256% (all plots) or 10–85% (thinned and burned plots). Thirteen percent of the pre‐settlement pines died in the treated area by 2003, but 9% percent also died in the control, indicating that pre‐settlement pines in untreated areas were nearly as vulnerable as those exposed to restoration treatments. Large snags increased 45%, and 65% of logs >50 cm were retained, achieving implementation goals. Although restoration treatments were not implemented totally to specifications, they were effective in attaining the overall project goal of restoring more open forest structure while preserving more than 75% of the pre‐settlement pines. Canopy fuel loads were substantially reduced, allowing for the reintroduction of surface fires.     

Historical Stem‐Mapped Permanent Plots Increase Precision of Reconstructed Reference Data in Ponderosa Pine Forests of Northern Arizona

Forest structural reference conditions are widely used to understand how ecosystems have been altered and guide restoration and management objectives. We used six stem‐mapped permanent plots established in the early twentieth century to provide precise structural reference conditions for ponderosa pine forests of northern Arizona prior to Euro‐American settlement. Reference conditions for these plots in 1873–1874 included the following historical attributes: tree densities of 45–127 trees/ha, mean tree diameter at breast height (dbh) of 43.8 cm with a corresponding quadratic mean diameter range of 41.5–51.3 cm, and a stand basal area of 9.2–18.0 m²/ha. The reconstructed diameter distributions (for live ponderosa pine trees with dbh ≥9.14 cm) prior to fire exclusion varied in shape but generally displayed an irregular unimodal distribution. We suggest that management objectives for the structural restoration of ponderosa pine forests of northern Arizona emphasize: (1) conservation and retention of all pre‐settlement (>130 years) trees; (2) reduction of tree densities with a restoration objective ranging between 50 and 150 trees/ha having a large‐tree component between 25 and 50% of the total trees per hectare, respectively; (3) manipulation of the diameter distribution to achieve a unimodal or irregular, uneven‐aged shape (possibly targeting a balanced, uneven‐aged shape on cinder soil types) through the use of harvest and thinning practices that mimic gap disturbances (i.e., individual tree selection system); and (4) retention of 3–11 snags and logs per hectare resulting from natural mortality.     

Does stand structure influence susceptibility of eucalypt floodplain forests to dieback?

Forest dieback is a worldwide problem that is likely to increase with climate change and increasing human demands for resources. Eucalyptus camaldulensis forests are an acute example of forest dieback, with 70% of the Victorian Murray River floodplain in some state of dieback. If we are to halt dieback in these floodplain forests, we need to understand what makes stands susceptible to dieback. Forest diebacks are often related to stand structure, with dieback more severe in senescent or high‐density stands. We determined whether certain stand structures make these forests more susceptible to dieback. We undertook an extensive survey of 176 stands across 100 000 ha of forest, covering the range of stand condition on this floodplain. Large and small trees (20‐, 40‐, 80‐ and 120‐cm diameter) showed a similar reduction in the probability of being alive with decreasing stand condition. A slight improvement in stand condition was found at higher densities and basal areas, which may reflect the higher productivity or younger age of these stands. Stand condition was moderately, positively correlated with longitude, with stand condition being higher in the east of the Murray River floodplain where flooding frequencies are currently higher. This suggests that dieback of these floodplain forests would be more effectively mitigated by increased water availability through flooding than by altering stand structure.     

A comparison of structural characteristics between old-growth and postfire second-growth hemlock–hardwood forests in Adirondack Park, New York, U. S. A.

1 This study compares the structural characteristics of 12 old‐growth and six postfire second‐growth hemlock–northern hardwood stands in north central Adirondack Park, New York, in order to test the null hypothesis that there are no differences in species composition, size structure, age structure and attributes such as dead wood and canopy gaps between old‐growth stands and this type of second‐growth forest. 2 The second‐growth forests of this study regenerated following widespread logging‐related fires in either 1903 or 1908; the old growth and second growth have similar environmental settings. 3 Estimates of stand ages, derived from an increment core of the oldest tree in each stand, range from 88 to 390 years. 4 Structural attributes are related to stand age (i.e. stage of development). In comparison with the second‐growth forests of this study, older stands are characterized as (a) a larger average diameter of canopy trees; (b) a greater basal area of trees; (c) a lower density of canopy trees and of all trees ≥ 10 cm d.b.h.; (d) a higher density of eastern hemlock (Tsuga canadensis (L.) Carrière) trees; (e) a higher density of large trees (≥ 50 cm d.b.h.); (f) larger canopy gaps; and (g) a greater volume of coarse woody debris (both logs ≥ 20 cm d.b.h. and snags ≥ 10 cm d.b.h.). 5 Despite differences between old growth and second growth, especially in species composition, it appears from observations of the 18 stands that second‐growth forests are developing some structural characteristics of old growth. 6 Structural attributes of the old‐growth forests are similar to characteristics of the same forest type in geographically distant areas in eastern USA.     

Gap formation and dynamics after long‐term steady state in an old‐growth Picea abies stand in Norway: Above‐ and belowground interactions

Stand dynamics and the gap initiation prior to gap formation are not well‐understood because of its long‐term nature and the scarcity of late‐successional stands. Reconstruction of such disturbance is normally based on historical records and dendroecological methods. We investigated gap initiation and formation at the fine‐scale stand level in the old‐growth reserve of Karlshaugen in Norway. Given its long‐term conservation history, and thorough mapping in permanent marked plots with spatially referenced trees, it provides an opportunity to present stand development before, during, and after gap formation. Late‐successional decline in biomass was recorded after more than 50 years of close to steady state. Gaps in the canopy were mainly created by large old trees that had been killed by spruce bark beetles. Snapping by wind was the main reason for treefall. Long‐term dominance of Norway spruce excluded downy birch and Scots pine from the stand. Comparisons of the forest floor soil properties between the gap and nongap area showed significantly higher concentrations of plant available Ca within the gap area. Plant root simulator (PRS?) probes showed significantly higher supply rates for Ca and Mg, but significantly lower K for the gap compared to the nongap area. Soil water from the gap area had significantly higher C:N ratios compared to the nongap area. Fine‐scale variation with increasing distance to logs indicated that CWD is important for leaking of DOC and Ca. Our long‐term study from Karlshaugen documents gap dynamics after more than 50 years of steady state and a multiscale disturbance regime in an old‐growth forest. The observed disturbance dynamic caused higher aboveground and belowground heterogeneity in plots, coarse woody debris, and nutrients. Our study of the nutrient levels of the forest floor suggest that natural gaps of old‐growth forest provide a long‐lasting biogeochemical feedback system particularly with respect to Ca and probably also N. Norway spruce trees near the gap edge responded with high plasticity to reduced competition, showing the importance of the edge zone as hot spots for establishing heterogeneity, but also the potential for carbon sequestration in old‐growth forest.     

Diversity and succession of epiphytic macrolichen communities in low‐elevation managed conifer forests in Western Oregon

Abstract. We examined epiphytic macrolichen communities in Pseudotsuga menziesii (Douglas‐fir) forests across the western Oregon landscape for relationships to environmental gradients, stand age and structure, and commercial thinning. We used a retrospective, blocked design through the Coast and the western Cascade ranges of Oregon. Each of our 17 blocks consisted of a young, unthinned stand (age 50–110 yr); an adjacent, thinned stand of equivalent age; and an old‐growth stand (age > 200 yr). We found 110 epiphytic macrolichen taxa in the stands. Forage‐providing alectorioid lichens and the nitrogen‐fixing cyanolichen Lobaria oregana associated strongly with old‐growth stands and remnant old trees in younger stands (unthinned + thinned). Relative to unthinned stands, thinned stands had a slightly higher abundance of alectorioid lichens and a greater presence of Hypogymnia imshaugii. However, thinned stands hosted a lower landscape‐level (γ) diversity, lacking many species that occurred infrequently in the unthinned stands. Patterns in the lichen community composition correlated strongly with climatic gradients; the greatest variation in composition was between the Coast and Cascade ranges. The difference in communities between mountain ranges was greatest among stands 70–110 yr old, suggesting a difference in lichen successional dynamics between the ranges.     

Modeling Ecological Restoration Effects on Ponderosa Pine Forest Structure

FIRESUM, an ecological process model incorporating surface fire disturbance, was modified for use in southwestern ponderosa pine ecosystems. The model was used to determine changes in forest structure over time and then applied to simulate changes in aboveground biomass and nitrogen storage since exclusion of the natural frequent fire regime in an unharvested Arizona forest. Dendroecological reconstruction of forest structure in 1876, prior to Euro‐American settlement, was used to initialize the model; projections were validated with forest measurements in 1992. Biomass allocations shifted from herbaceous plants to trees, and nitrogen was increasingly retained in living and dead tree biomass over the 116‐year period (1876–1992). Forest conditions in 1992 were substantially degraded compared to reference presettlement conditions: old‐growth trees were dying at accelerated rates, herbaceous production was reduced nearly 90%, and the entire stand was highly susceptible to high‐intensity wildfire. Following an experiment initiated in 1993 to test ecological restoration treatments, future changes were modeled for the next century. Future forest structure remained within the natural presettlement range of variability under the full restoration treatment, in which forest biomass structure was thinned to emulate presettlement conditions and repeated low‐intensity fire was reintroduced. Simulation of the control treatment indicated continuation of exceptionally high tree density, probably culminating in stand‐replacing ecosystem change through high‐intensity wildfire or tree mortality from pathogens. Intermediate results were observed in the partial restoration treatment (tree thinning only); the open forest structure and high herbaceous productivity found immediately after treatment were gradually degraded as dense tree cover reestablished in the absence of fire. Modeling results support comprehensive restorative management as a long‐term approach to conservation of key indigenous ecosystem characteristics.