Community occupancy responses of small mammals to restoration treatments in ponderosa pine forests, northern Arizona, USA

In western North American conifer forests, wildfires are increasing in frequency and severity due to heavy fuel loads that have accumulated after a century of fire suppression. Forest restoration treatments (e.g., thinning and/or burning) are being designed and implemented at large spatial and temporal scales in an effort to reduce fire risk and restore forest structure and function. In ponderosa pine (Pinus ponderosa) forests, predominantly open forest structure and a frequent, low-severity fire regime constituted the evolutionary environment for wildlife that persisted for thousands of years. Small mammals are important in forest ecosystems as prey and in affecting primary production and decomposition. During 2006-2009, we trapped eight species of small mammals at 294 sites in northern Arizona and used occupancy modeling to determine community responses to thinning and habitat features. The most important covariates in predicting small mammal occupancy were understory vegetation cover, large snags, and treatment. Our analysis identified two generalist species found at relatively high occupancy rates across all sites, four open-forest species that responded positively to treatment, and two dense-forest species that responded negatively to treatment unless specific habitat features were retained. Our results indicate that all eight small mammal species can benefit from restoration treatments, particularly if aspects of their evolutionary environment (e.g., large trees, snags, woody debris) are restored. The occupancy modeling approach we used resulted in precise species-level estimates of occupancy in response to habitat attributes for a greater number of small mammal species than in other comparable studies. We recommend our approach for other studies faced with high variability and broad spatial and temporal scales in assessing impacts of treatments or habitat alteration on wildlife species. Moreover, since forest planning efforts are increasingly focusing on progressively larger treatment implementation, better and more efficiently obtained ecological information is needed to inform these efforts.     

Demography of northern flying squirrels informs ecosystem management of western interior forests.

We studied northern flying squirrel (Glaucomys sabrinus) demography in the eastern Washington Cascade Range to test hypotheses about regional and local abundance patterns and to inform managers of the possible effects of fire and fuels management on flying squirrels. We quantified habitat characteristics and squirrel density, population trends, and demography in three typical forest cover types over a four-year period. We had 2034 captures of flying squirrels over 41 000 trap nights from 1997 through 2000 and marked 879 squirrels for mark-recapture population analysis. Ponderosa pine (Pinus ponderosa) forest appeared to be poorer habitat for flying squirrels than young or mature mixed-conifer forest. About 35% fewer individuals were captured in open pine forest than in dry mixed-conifer Douglas-fir (Pseudotsuga menziesii) and grand fir (Abies grandis) forests. Home ranges were 85% larger in pine forest (4.6 ha) than in mixed-conifer forests (2.5 ha). Similarly, population density (Huggins estimator) in ponderosa pine forest was half (1.1 squirrels/ha) that of mixed-conifer forest (2.2 squirrels/ha). Tree canopy cover was the single best correlate of squirrel density (r = 0.77), with an apparent threshold of 55% canopy cover separating stands with low- from high-density populations. Pradel estimates of annual recruitment were lower in open pine (0.28) than in young (0.35) and mature (0.37) forest. High recruitment was most strongly associated with high understory plant species richness and truffle biomass. Annual survival rates ranged from 45% to 59% and did not vary among cover types. Survival was most strongly associated with understory species richness and forage lichen biomass. Maximum snow depth had a strong negative effect on survival. Rate of per capita increase showed a density-dependent response. Thinning and prescribed burning in ponderosa pine and dry mixed conifer forests to restore stable fire regimes and forest structure might reduce flying squirrel densities at stand levels by reducing forest canopy, woody debris, and the diversity or biomass of understory plants, truffles, and lichens. Those impacts might be ameliorated by patchy harvesting and the retention of large trees, woody debris, and mistletoe brooms. Negative stand-level impacts would be traded for increased resistance and resilience of dry-forest landscapes to now-common, large-scale stand replacement fires.     

Frequent fire alters nitrogen transformations in ponderosa pine stands of the inland northwest

Recurrent, low-severity fire in ponderosa pine (Pinus ponderosa)/interior Douglas-fir (Pseudotsuga menziesii var. glauca) forests is thought to have directly influenced nitrogen (N) cycling and availability. However, no studies to date have investigated the influence of natural fire intervals on soil processes in undisturbed forests, thereby limiting our ability to understand ecological processes and successional dynamics in this important ecosystem of the Rocky Mountain West. Here, we tested the standing hypothesis that recurrent fire in ponderosa pine/Douglas-fir forests of the Inland Northwest decreases total soil N, but increases N turnover and nutrient availability. We compared soils in stands unburned over the past 69-130 years vs. stands exposed to two or more fires over the last 130 years at seven distinct locations in two wilderness areas. Mineral soil samples were collected from each of the seven sites in June and July of 2003 and analyzed for pH, total C and N, potentially mineralizable N (PMN), and extractable NH4+, NO3-, PO4(-3), Ca+2, Mg+2, and K+. Nitrogen transformations were assessed at five sites by installing ionic resin capsules in the mineral soil in August of 2003 and by conducting laboratory assays of nitrification potential and net nitrification in aerobic incubations. Total N and PMN decreased in stands subjected to multiple fires. This loss of total N and labile N was not reflected in concentrations of extractable NH4+ and NO3-. Rather, multiple fires caused an increase in NO3 sorbed on ionic resins, nitrification potential, and net nitrification in spite of the burned stands not having been exposed to fire for at least 12-17 years. Charcoal collected from a recent fire site and added to unburned soils increased nitrification potential, suggesting that the decrease of charcoal in the absence of fire may play an important role in N transformations in fire-dependent ecosystems in the long term. Interestingly, we found no consistent effect of fire frequency on extractable P or alkaline metal concentrations. Our results corroborate the largely untested hypothesis that frequent fire in ponderosa pine forests increases inorganic N availability in the long term and emphasize the need to study natural, unmanaged sites in far greater detail.     

Homeostatic maintenance of ponderosa pine gas exchange in response to stand density changes.

Homeostatic maintenance of gas exchange optimizes carbon gain per water loss. Homeostasis is regulated by short-term physiological and long-term structural mechanisms, both of which may respond to changes in resource availability associated with competition. Therefore, stand density regulation via silvicultural manipulations may facilitate growth and survival through mechanisms operating at both short and long timescales. We investigated the responses of ponderosa pine (Pinus ponderosa) to stand basal area manipulations in Arizona, USA. Stand basal area was manipulated to seven replicated levels in 1962 and was maintained for four decades by decadal thinning. We measured basal area increment (BAI) to assess the response and sustainability of wood growth, carbon isotope discrimination (A) inferred from annual rings to assess the response of crown gas exchange, and ratios of leaf area to sapwood area (A(l):A(s)) to assess longer term structural acclimation. Basal area treatments increased soil water potential (r2 = 0.99) but did not affect photosynthetic capacity. BAI increased within two years of thinning, and the 40-year mean BAI was negatively correlated with stand basal area (r2 = 0.98). delta was negatively correlated with stand basal area for years 5 through 12 after thinning (r2 = 0.90). However, delta was relatively invariant with basal area for the period 13-40 years after initial thinning despite maintenance of treatment basal areas via repeated decadal thinnings. Independent gas exchange measurements verified that the ratio of photosynthesis to stomatal conductance was invariant with basal area, but absolute values of both were elevated at lower basal areas. A(l):A(s) was negatively correlated with basal area (r2 = 0.93). We hypothesize that increased A(l):A(s) is a homeostatic response to increased water availability that maximizes water-use efficiency and whole-tree carbon uptake. Elevated A(l):A(s) of trees at low basal areas was associated with greater resilience to climate, i.e., greater absolute BAI during drought; however, trees with high A(l):A(s) in low basal area stands also exhibited the greatest sensitivity to drought, i.e., greater relative decline in BAI.     

Nitrogen spatial heterogeneity influences diversity following restoration in a ponderosa pine forest, Montana.

The resource heterogeneity hypothesis (RHH) is frequently cited in the ecological literature as an important mechanism for maintaining species diversity. The RHH has rarely been evaluated in the context of restoration ecology in which a commonly cited goal is to restore diversity. In this study we focused on the spatial heterogeneity of total inorganic nitrogen (TIN) following restoration treatments in a ponderosa pine (Pinus ponderosa)/Douglas-fir (Pseudotsuga menziesii) forest in western Montana, USA. Our objective was to evaluate relationships between understory species richness and TIN heterogeneity following mechanical thinning (thin-only), prescribed burning (burn-only), and mechanical thinning with prescribed burning (thin/burn) to discern the ecological and management implications of these restoration approaches. We employed a randomized block design, with three 9-ha replicates of each treatment and an untreated control. Within each treatment, we randomly established a 20 x 50 m (1000 m2) plot in which we measured species richness across the entire plot and in 12 1-m(2) quadrats randomly placed within each larger plot. Additionally, we measured TIN from a grid consisting of 112 soil samples (0-5 cm) in each plot and computed standard deviations as a measure of heterogeneity. We found a correlation between the net increase in species richness and the TIN standard deviations one and two years following restoration treatments, supporting RHH. Using nonmetric multidimensional scaling ordination and chi-squared analysis, we found that high and low TIN quadrats contained different understory communities in 2003 and 2004, further supporting RHH. A comparison of restoration treatments demonstrated that thin/burn and burn-only treatments created higher N heterogeneity relative to the control. We also found that within prescribed burn treatments, TIN heterogeneity was positively correlated with fine-fuel consumption, a variable reflecting burn severity. These findings may lead to more informed restoration decisions that consider treatment effects on understory diversity in ponderosa pine/Douglas-fir ecosystems.     

Simulation of wildfire effects on the nitrogen cycle of a Pinus banksiana ecosystem in New Brunswick, Canada

A non-linear, deterministic model of biomass accumulation and nitrogen cycling in an even-aged, pure jack pine (Pinus banksiana Lamb.) stand was developed and used to explore effects of fire intensity and frequency of burning on the long-term nitrogen cycle. Given the model structure and assumptions, simulated results showed that successive fires at both light and severe fire intensities caused gradual depletion of the amount of N accumulated in the vegetation layers. Fires also reduced the amount of N in the litter and soil pools, with the initially large soil organically-bound N pool showing a particularly sharp decline, and decreased the productivity of the simulated stand. A frequency of one fire per 20 years for five successive burns produced declines of N accumulated in the tree stratum of 50–75% (depending upon fire intensity) in comparison with the undisturbed system at a corresponding age, whereas a 100-year frequency produced decreases of 10–22%. Similarly, declines in litter layer N were 54–72% at a 40-year frequency, compared with 30–55% at a 100-year frequency. The simulated results also suggested that both the stand age when burning occurred and the fire frequency were important, because distinctive patterns of accumulation and decline of N in ecosystem pools existed with increasing stand age. A serious lack of information regarding processes inherent in the model was found to exist in certain cases. Important processes which are currently poorly quantified include: (1) the factors controlling rates of tree growth; (2) the relation of foliar and other tissue N to soil N concentrations and foliar translocation; (3) the relation of forest floor conditions to decomposition and stand structural characteristics; and (4) the controls of a variety of soil N transformations, transfers, leaching and decomposition rates. Because of this basic lack of information and the great dependence of the model's behavior on these processes, the present version of the model is not suitable for real-world prediction. The model does have use as a means of combining hypotheses about a system into an explicit structure and examining the collective consequences of this, as well as pointing out future research needs for the system.     

Carbon stocks across a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands

Forests function as a major global C sink, and forest management strategies that maximize C stocks offer one possible means of mitigating the impacts of increasing anthropogenic CO2 emissions. We studied the effects of thinning, a common management technique in many forest types, on age-related trends in C stocks using a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands ranging from 9 to 306 years old. Live tree C stocks increased with age to a maximum near the middle of the chronosequence in unmanaged stands, and increased across the entire chronosequence in thinned stands. C in live understory vegetation and C in the mineral soil each declined rapidly with age in young stands but changed relatively little in middle-aged to older stands regardless of management. Forest floor C stocks increased with age in unmanaged stands, but forest floor C decreased with age after the onset of thinning around age 40 in thinned stands. Deadwood C was highly variable, but decreased with age in thinned stands. Total ecosystem C increased with stand age until approaching an asymptote around age 150. The increase in total ecosystem C was paralleled by an age-related increase in total aboveground C, but relatively little change in total belowground C. Thinning had surprisingly little impact on total ecosystem C stocks, but it did modestly alter age-related trends in total ecosystem C allocation between aboveground and belowground pools. In addition to characterizing the subtle differences in C dynamics between thinned and unmanaged stands, these results suggest that C accrual in red pine stands continues well beyond the 60-100 year management rotations typical for this system. Management plans that incorporate longer rotations and thinning in some stands could play an important role in maximizing C stocks in red pine forests while meeting other objectives including timber extraction, biodiversity conservation, restoration, and fuel reduction goals.     

离子交换树脂袋法研究森林土壤硝态氮及其对氮沉降增加的响应

用离子交换树脂袋法，研究了鼎湖山三种森林（马尾松林、马尾松针叶阔叶混交林和季风常绿阔叶林）土壤硝态氮对外加氮的响应特征。结果表明，土壤硝态氮显著地受森林类型、季节和氮处理的影响。整体而言，阔叶林土壤硝态氮极显著高于马尾松林和混交林，而马尾松林和混交林之间的差异则不显著。三种森林土壤硝态氮的季节变化均表现为春季和夏季极显著高于冬季和秋季，而冬季又显著高于秋季。外加氮处理提高土壤硝态氮水平，其中在马尾松林和阔叶林氮处理效应显著。所得结果与直接采集土壤或土壤水测定的硝态氮含量的结果一致，表明离子交换树脂袋法是评价土壤硝态氮水平行之有效的手段之一。     

Litter and dead wood dynamics in ponderosa pine forests along a 160-year chronosequence.

Disturbances such as fire play a key role in controlling ecosystem structure. In fire-prone forests, organic detritus comprises a large pool of carbon and can control the frequency and intensity of fire. The ponderosa pine forests of the Colorado Front Range, USA, where fire has been suppressed for a century, provide an ideal system for studying the long-term dynamics of detrital pools. Our objectives were (1) to quantify the long-term temporal dynamics of detrital pools; and (2) to determine to what extent present stand structure, topography, and soils constrain these dynamics. We collected data on downed dead wood, litter, duff (partially decomposed litter on the forest floor), stand structure, topographic position, and soils for 31 sites along a 160-year chronosequence. We developed a compartment model and parameterized it to describe the temporal trends in the detrital pools. We then developed four sets of statistical models, quantifying the hypothesized relationship between pool size and (1) stand structure, (2) topography, (3) soils variables, and (4) time since fire. We contrasted how much support each hypothesis had in the data using Akaike's Information Criterion (AIC). Time since fire explained 39-80% of the variability in dead wood of different size classes. Pool size increased to a peak as material killed by the fire fell, then decomposed rapidly to a minimum (61-85 years after fire for the different pools). It then increased, presumably as new detritus was produced by the regenerating stand. Litter was most strongly related to canopy cover (r2 = 77%), suggesting that litter fall, rather than decomposition, controls its dynamics. The temporal dynamics of duff were the hardest to predict. Detrital pool sizes were more strongly related to time since fire than to environmental variables. Woody debris peak-to-minimum time was 46-67 years, overlapping the range of historical fire return intervals (1 to > 100 years). Fires may therefore have burned under a wide range of fuel conditions, supporting the hypothesis that this region's fire regime was mixed severity.     

东北过伐林区不同林分类型土壤肥力质量评价研究

保持和提高森林土壤肥力质量是实现森林健康的基础。研究不同林型下土壤肥力质量对森林可持续经营具有重要意义。以东北过伐林区金仓林场中的落叶松Lartx gmelinii人工纯林、云杉Picea asperatax紫椴Tilia amurensis天然混交林、紫椴×白桦Betula platyphylla天然混交林、五角槭Acer monox白桦×落叶松天然混交林、五角槭×白桦×紫椴天然混交林、白桦×落叶松天然混交林和五角械×紫椴×青杨Populus pseudo-simonii天然混交林为研究对象,分析和比较了不同林分下的土壤物理和化学性质,并采用主成分分析与聚类分析相结合的方法评价了其土壤肥力质量。结果表明：①林分类型对土壤理化性质影响显著;随土壤深度增加,土壤密度和土壤pH值增大,而土壤含水量、物理性粘粒含量和养分质量分数减少,但其在不同林分下的变化程度不同;②土壤因子之间关系密切,物理性粘粒含量与含水量、全氮、速效钾质量分数呈极显著正相关,而与土壤密度和pH值呈显著负相关;土壤养分质量分数之间具有不同程度的显著正相关关系;③采用主成分分析法对不同林分下0～60cm的土壤肥力质量进行了评价,土壤肥力质量表现为天然混交林高于落叶松人工纯林;采伐降低了土壤的肥力质量;在落叶松人工纯林中,随着林龄的增加,土壤的肥力质量下降。通过对土壤肥力质量综合指标值的聚类分析,将研究区土壤肥力质量分为优、良、中、差4个等级,肥力质量属于中等以上（优、良、中等）的林地所占研究样地的比例为66．7％,研究区森林土壤肥力质量属于中等水平。建议在经营过伐林时,注意调整林分结构和树种组成,控制采伐强度,同时进行长期定位观测和比较,以改善林分整体的肥力状况。本研究结果为该地区林业可持续经营提供直接依据,也为东北地区森林土壤肥力质量评价提供参考。     

Agricultural legacies in forest environments: tree communities, soil properties, and light availability.

Temperate deciduous forests across much of Europe and eastern North America reflect legacies of past land use, particularly in the diversity and composition of plant communities. Intense disturbances, such as clearing forests for agriculture, may cause persistent environmental changes that continue to shape vegetation patterns as landscapes recover. We assessed the long-term consequences of agriculture for environmental conditions in central New York forests, including tree community structure and composition, soil physical and chemical properties, and light availability. To isolate the effects of agriculture, we compared 20 adjacent pairs of forests that were never cleared for agriculture (primary forests) and forests that established 85-100 years ago on plowed fields (secondary forests). Tree communities in primary and secondary forests had similar stem density, though secondary forests had 14% greater basal area. Species composition differed dramatically between the two forest types, with primary forests dominated by Acer saccharum and Fagus grandifolia and secondary forests by Acer rubrum and Pinus strobus. Primary and secondary forests showed no consistent differences in soil physical properties or in the principal gradient of soil fertility associated with soil pH. Within stands, however, soil water content and pH were more variable in primary forests. Secondary forest soils had 15% less organic matter, 16% less total carbon, and 29% less extractable phosphorus in the top 10 cm than adjacent primary stands, though the ranges of the forest types mostly overlapped. Understory light availability in primary and secondary forests was similar. These results suggest that, within 100 years, post-agricultural stands have recovered conditions comparable to less disturbed forests in many attributes, including tree size and number, soil physical properties, soil chemical properties associated with pH, and understory light availability. The principal legacies of agriculture that remain in these forests are the reduced levels of soil organic matter, carbon, and phosphorus; the spatial homogenization of soil properties; and the altered species composition of the vegetation.     

Atmospheric deposition may affect northern hardwood forest composition by altering soil nutrient supply.

In the northeastern United States, the input of reactive nitrogen (N) via atmospheric deposition has increased rapidly since the onset of the industrial revolution. During the same period of time, acid precipitation and forest harvest have removed substantial quantities of base cations from soil. Because of the dominance of base-poor soils and the low rates of atmospheric base cation deposition, soils throughout the northeastern United States may be increasingly rich in N but poor in calcium (Ca). We studied the consequences of a change in soil N and Ca availability on forest composition by transplanting seedlings of four tree species into replicate plots in the understory and in canopy gaps amended with N and Ca in factorial combination. In this paper, we report on the growth and survivorship of seedlings over a four-year period. Relative to control plots, fertilization with N increased red maple growth by an average of 39% whereas fertilization with Ca decreased survivorship in the understory by 41%. In sugar maple, fertilization with Ca increased growth by 232% and 46% in the forest understory and in canopy gaps, respectively, and significantly increased high light survivorship. Fertilization with N decreased white pine survivorship by 69% in the understory whereas high Ca availability significantly increased survivorship. Fertilization with N or Ca alone reduced red oak growth but had no effect on survivorship. The results of this study suggest that historical losses of soil Ca and the continuing effects of atmospheric-N deposition on N availability are likely to alter the composition of northeastern North American forests because of the positive effects of N enrichment on the growth of red maple and the negative effects of Ca loss on the growth and survivorship of sugar maple and white pine.     

Calcium constrains plant control over forest ecosystem nitrogen cycling

Forest ecosystem nitrogen (N) cycling is a critical controller of the ability of forests to prevent the movement of reactive N to receiving waters and the atmosphere and to sequester elevated levels of atmospheric carbon dioxide (CO2). Here we show that calcium (Ca) constrains the ability of northern hardwood forest trees to control the availability and loss of nitrogen. We evaluated soil N-cycling response to Ca additions in the presence and absence of plants and observed that when plants were present, Ca additions "tightened" the ecosystem N cycle, with decreases in inorganic N levels, potential net N mineralization rates, microbial biomass N content, and denitrification potential. In the absence of plants, Ca additions induced marked increases in nitrification (the key process controlling ecosystem N losses) and inorganic N levels. The observed "tightening" of the N cycle when Ca was added in the presence of plants suggests that the capacity of forests to absorb elevated levels of atmospheric N and CO2 is fundamentally constrained by base cations, which have been depleted in many areas of the globe by acid rain and forest harvesting.     

Changes in microbial community characteristics and soil organic matter with nitrogen additions in two tropical forests

Microbial communities and their associated enzyme activities affect the amount and chemical quality of carbon (C) in soils. Increasing nitrogen (N) deposition, particularly in N-rich tropical forests, is likely to change the composition and behavior of microbial communities and feed back on ecosystem structure and function. This study presents a novel assessment of mechanistic links between microbial responses to N deposition and shifts in soil organic matter (SOM) quality and quantity. We used phospholipid fatty acid (PLFA) analysis and microbial enzyme assays in soils to assess microbial community responses to long-term N additions in two distinct tropical rain forests. We used soil density fractionation and 13C nuclear magnetic resonance (NMR) spectroscopy to measure related changes in SOM pool sizes and chemical quality. Microbial biomass increased in response to N fertilization in both tropical forests and corresponded to declines in pools of low-density SOM. The chemical quality of this soil C pool reflected ecosystem-specific changes in microbial community composition. In the lower-elevation forest, there was an increase in gram-negative bacteria PLFA biomass, and there were significant losses of labile C chemical groups (O-alkyls). In contrast, the upper-elevation tropical forest had an increase in fungal PLFAs with N additions and declines in C groups associated with increased soil C storage (alkyls). The dynamics of microbial enzymatic activities with N addition provided a functional link between changes in microbial community structure and SOM chemistry. Ecosystem-specific changes in microbial community composition are likely to have far-reaching effects on soil carbon storage and cycling. This study indicates that microbial communities in N-rich tropical forests can be sensitive to added N, but we can expect significant variability in how ecosystem structure and function respond to N deposition among tropical forest types.     

降水变化、氮添加对鼎湖山主要森林土壤有机碳矿化和土壤微生物碳的影响

全球变化对土壤有机碳(SOC)存贮与分解的影响在全球碳(C)循环中具有重要地位.分别通过室内土壤培养法和氯仿熏蒸法,研究了降水变化和氮(N)添加处理对鼎湖山3种不同演替阶段的季风常绿阔叶林、针阔混交林和马尾松针叶林SOC矿化和土壤微生物量碳(SMBC)的影响.结果表明:1)降水量增加能够提高森林演替晚期SOC累积矿化量和矿化速率,而对森林演替早期SOC累积矿化量和矿化速率没有显著影响(P>0.05).2)干旱条件(降水量减少)降低森林SMBC含量,且在鼎湖山季风林表层土壤(0～10 cm)中SMBC的减少达到显著水平(P<0.05).3)N添加处理对鼎湖山3种森林类型SOC累积矿化量、矿化速率以及SMBC都没有显著影响(P>0.05).未来关于SOC矿化对全球变化响应的研究,要综合考虑土壤有机质质量、C/N比例、外源性氮输入等因素的作用.图4表2参37     

Thinning and burning result in low-level invasion by nonnative plants but neutral effects on natives.

Many historically fire-adapted forests are now highly susceptible to damage from insects, pathogens, and stand-replacing fires. As a result, managers are employing treatments to reduce fuel loadings and to restore the structure, species, and processes that characterized these forests prior to widespread fire suppression, logging, and grazing. However, the consequences of these activities for understory plant communities are not well understood. We examined the effects of thinning and prescribed fire on plant composition and diversity in Pinus ponderosa forests of eastern Washington (USA). Data on abundance and richness of native and nonnative plants were collected in 70 stands in the Colville, Okanogan, and Wenatchee National Forests. Stands represented one of four treatments: thinning, burning, thinning followed by burning, or control; treatments had been conducted 3-19 years before sampling. Multi-response permutation procedures revealed no significant effect of thinning or burning on understory plant composition. Similarly, there were no significant differences among treatments in cover or richness of native plants. In contrast, nonnative plants showed small, but highly significant, increases in cover and richness in response to both thinning and burning. In the combined treatment, cover of nonnative plants averaged 2% (5% of total plant cover) but did not exceed 7% (16% of total cover) at any site. Cover and richness of nonnative herbs showed small increases with intensity of disturbance and time since treatment. Nonnative plants were significantly less abundant in treated stands than on adjacent roadsides or skid trails, and cover within these potential source areas explained little of the variation in abundance within treated stands. Although thinning and burning may promote invasion of nonnative plants in these forests, our data suggest that their abundance is limited and relatively stable on most sites.     

持续提高黄土高原植被水土保持功能的配套技术(I)森林保持水土的条件

阐述了森林保持水土的条件 ,即良好的林分结构、必要的面积比例和一定的林龄 ,揭示了成林过程中水土保持功能的动态变化 ,为研究防护林体系水土保持功能持续提高技术提供了理论基础     

凋落物处理对森林地表CO2通量的影响及其调控机理

全球变化和森林演替可以导致森林地表凋落物数量和质量的变化,从而对森林地表 CO2通量产生影响。本实验对亚热带不同演替阶段的3种,即马尾松林（前期）、混交林（中期）和季风林（后期）进行地表凋落物去除、加倍与置换处理,利用静态箱气相色谱法测定地表CO2通量,并同步测定气温、土壤温度和湿度,分析凋落物质量和数量变化对森林地表CO2通量的影响及其调控机理。结果表明,（1）去除凋落物处理显著降低了不同演替阶段的3种森林地表CO2通量,而加倍凋落物处理可以增加森林地表CO2通量,但不同演替阶段增加的幅度不同,依次为：季风林＞马尾松林＞混交林。（2）置换凋落物对不同演替阶段的森林地表 CO2通量的影响不同,在演替后期的季风林中,置换混交林和马尾松林凋落物处理均增加地表CO2通量;在演替中期的混交林中,置换季风林和马尾松林凋落物均降低地表CO2通量。在演替前期的马尾松林中,置换季风林凋落物增加地表CO2通量,而置换混交林凋落物降低了地表CO2通量（3）结合测定的土壤温度和水分数据分析得出,凋落物处理引起森林地表 CO2通量的变化是通过处理凋落物质量和数量后改变森林地表水热条件来实现的。（4）3个林型的各种处理,地表 CO2通量与土壤温度均呈显著的指数相关关系,但不同处理不同地改变了森林地表土壤 CO2通量对温度的敏感性,即Q10值。     

Climate effects on fire regimes and tree recruitment in Black Hills ponderosa pine forests

Climate influences forest structure through effects on both species demography (recruitment and mortality) and disturbance regimes. Here, I compare multi-century chronologies of regional fire years and tree recruitment from ponderosa pine forests in the Black Hills of southwestern South Dakota and northeastern Wyoming to reconstructions of precipitation and global circulation indices. Regional fire years were affected by droughts and variations in both Pacific and Atlantic sea surface temperatures. Fires were synchronous with La Ni?as, cool phases of the Pacific Decadal Oscillation (PDO), and warm phases of the Atlantic Multidecadal Oscillation (AMO). These quasi-periodic circulation features are associated with drought conditions over much of the western United States. The opposite pattern (El Ni?o, warm PDO, cool AMO) was associated with fewer fires than expected. Regional tree recruitment largely occurred during wet periods in precipitation reconstructions, with the most abundant recruitment coeval with an extended pluvial from the late 1700s to early 1800s. Widespread even-aged cohorts likely were not the result of large crown fires causing overstory mortality, but rather were caused by optimal climate conditions that contributed to synchronous regional recruitment and longer intervals between surface fires. Synchronous recruitment driven by climate is an example of the Moran effect. The presence of abundant fire-scarred trees in multi-aged stands supports a prevailing historical model for ponderosa pine forests in which recurrent surface fires affected heterogenous forest structure, although the Black Hills apparently had a greater range of fire behavior and resulting forest structure over multi-decadal time scales than ponderosa pine forests of the Southwest that burned more often.     

不同林龄马尾松（Pinus massoniana）人工林碳氮磷分配格局及化学计量特征

为了解长江上游低山丘陵区马尾松（Pinus massoniana）人工林生态系统的C、N、P分配格局及化学计量特征,本文采用时空互代的方法,在宜宾高县来复林区选取三种不同林龄（5年生幼龄林、14年生中龄林、39年生成熟林）,但立地条件相近、样地情况基本一致的马尾松（Pinus massoniana）人工林作为研究对象,对马尾松针叶、凋落物及土壤中的C、N、P含量及 w（C）？w（N）？w（P）化学计量特征进行测定和分析。结果表明,（1）C、N、P 含量均表现为针叶〉凋落物〉土壤,且在三个库之间差异显著;（2）林龄对针叶、凋落物、土壤的 C、N、P 及 w（C）？w（N）、w（C）？w（P）计量比均有显著影响。（3）土壤 C、N、P含量在成熟林中最高;针叶和凋落物的C含量在成熟林中最低,N、P含量则在中龄林中最高。（4）随林龄增加马尾松对N、P的利用效率降低,针叶、凋落物及土壤的w（C）？w（N）与 w（C）？w（P）均表现为下降。（5）马尾松针叶w（N）？w（P）比值在14.37~15.53之间,说明该地区马尾松人工林受N和P的共同限制,但林龄对N、P养分限制的影响不显著。为提高该区马尾松人工林的生产力,建议在人工林的抚育管理中要适当增加N肥和P肥,同时也可在马尾松人工林引入豆科固氮植物以提高地力。该研究将马尾松针叶、凋落物及土壤结合起来探究随林龄增长C、N、P养分元素的分配格局及化学计量特征的变化,有助于全面、系统地揭示马尾松人工林生态系统的养分循环,对指导马尾松人工林生产,调节和改善林木生长环境,提高系统的养分利用效率及林地生产力具有重要意义。