Understory vegetation response to mountain pine beetle disturbance in northern Colorado lodgepole pine forests

Understory plants are an important element of forests, having a considerable influence on ecosystem functioning and canopy-tree development following disturbance. Recent bark beetle outbreaks across western North American forests have caused extensive canopy mortality, creating new growing conditions that provide the opportunity for changes within the intact understory. Over a five-year period following peak mountain pine beetle (MPB) activity across lodgepole pine-dominated forests in Rocky Mountain National Park, Colorado, we measured the changes in plant diversity, cover, and dominance by lifeform and quantified tree regeneration rates. Average species richness and diversity increased, but overall plant cover did not change. Graminoids appeared to benefit the most, increasing in average cover, richness, and relative dominance. The rise in graminoid dominance was largely at the expense of shrubs, which showed little ability to benefit from overstory mortality within the first years following attack. Most plant responses were positively related to the total tree basal area lost since the peak of the outbreak, suggesting that increased resource availability following tree death may benefit understory plants. However, a negative relationship between several understory variables and tree sapling density provides evidence that understory plants compete with saplings for the newly available resources. Tree seedling density nearly doubled over the duration of the study, indicating a strong regeneration pulse. Among species, lodgepole pine displayed the greatest tree seedling establishment. This is one of the first studies to use repeated measurements to describe this often-overlooked component of forest change associated with MPB disturbance.

     

Effects of Dwarf Mistletoe on Stand Structure of Lodgepole Pine Forests 21-28 Years Post-Mountain Pine Beetle Epidemic in Central Oregon

Lodgepole pine (Pinus contorta) forests are widely distributed throughout North America and are subject to mountain pine beetle (Dendroctonus ponderosae) epidemics, which have caused mortality over millions of hectares of mature trees in recent decades. Mountain pine beetle is known to influence stand structure, and has the ability to impact many forest processes. Dwarf mistletoe (Arceuthobium americanum) also influences stand structure and occurs frequently in post-mountain pine beetle epidemic lodgepole pine forests. Few studies have incorporated both disturbances simultaneously although they co-occur frequently on the landscape. The aim of this study is to investigate the stand structure of lodgepole pine forests 21–28 years after a mountain pine beetle epidemic with varying levels of dwarf mistletoe infection in the Deschutes National Forest in central Oregon. We compared stand density, stand basal area, canopy volume, proportion of the stand in dominant/codominant, intermediate, and suppressed cohorts, average height and average diameter of each cohort, across the range of dwarf mistletoe ratings to address differences in stand structure. We found strong evidence of a decrease in canopy volume, suppressed cohort height, and dominant/codominant cohort diameter with increasing stand-level dwarf mistletoe rating. There was strong evidence that as dwarf mistletoe rating increases, proportion of the stand in the dominant/codominant cohort decreases while proportion of the stand in the suppressed cohort increases. Structural differences associated with variable dwarf mistletoe severity create heterogeneity in this forest type and may have a significant influence on stand productivity and the resistance and resilience of these stands to future biotic and abiotic disturbances. Our findings show that it is imperative to incorporate dwarf mistletoe when studying stand productivity and ecosystem recovery processes in lodgepole pine forests because of its potential to influence stand structure.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Changes to the N cycle following bark beetle outbreaks in two contrasting conifer forest types

Outbreaks of Dendroctonus beetles are causing extensive mortality in conifer forests throughout North America. However, nitrogen (N) cycling impacts among forest types are not well known. We quantified beetle-induced changes in forest structure, soil temperature, and N cycling in Douglas-fir (Pseudotsuga menziesii) forests of Greater Yellowstone (WY, USA), and compared them to published lodgepole pine (Pinus contorta var. latifolia) data. Five undisturbed stands were compared to five beetle-killed stands (4–5 years post-outbreak). We hypothesized greater N cycling responses in Douglas-fir due to higher overall N stocks. Undisturbed Douglas-fir stands had greater litter N pools, soil N, and net N mineralization than lodgepole pine. Several responses to disturbance were similar between forest types, including a pulse of N-enriched litter, doubling of soil N availability, 30–50 % increase in understory cover, and 20 % increase in foliar N concentration of unattacked trees. However, the response of some ecosystem properties notably varied by host forest type. Soil temperature was unaffected in Douglas-fir, but lowered in lodgepole pine. Fresh foliar %N was uncorrelated with net N mineralization in Douglas-fir, but positively correlated in lodgepole pine. Though soil ammonium and nitrate, net N mineralization, and net nitrification all doubled, they remained low in both forest types (<6 μg N g soil^?1 NH₄ ⁺or NO₃ ^?; <25 μg N g soil^?1 year^?1 net N mineralization; <8 μg N g soil^?1 year^?1 net nitrification). Results suggest that beetle disturbance affected litter and soil N cycling similarly in each forest type, despite substantial differences in pre-disturbance biogeochemistry. In contrast, soil temperature and soil N–foliar N linkages differed between host forest types. This result suggests that disturbance type may be a better predictor of litter and soil N responses than forest type due to similar disturbance mechanisms and disturbance legacies across both host–beetle systems.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

Patterns of Biomass and Carbon Distribution across a Chronosequence of Chinese Pine (Pinus tabulaeformis) Forests

Patterns of biomass and carbon (C) storage distribution across Chinese pine (Pinus tabulaeformis) natural secondary forests are poorly documented. The objectives of this study were to examine the biomass and C pools of the major ecosystem components in a replicated age sequence of P. tabulaeformis secondary forest stands in Northern China. Within each stand, biomass of above- and belowground tree, understory (shrub and herb), and forest floor were determined from plot-level investigation and destructive sampling. Allometric equations using the diameter at breast height (DBH) were developed to quantify plant biomass. C stocks in the tree and understory biomass, forest floor, and mineral soil (0–100 cm) were estimated by analyzing the C concentration of each component. The results showed that the tree biomass of P. tabulaeformis stands was ranged from 123.8 Mg·ha^–1 for the young stand to 344.8 Mg·ha^–1 for the mature stand. The understory biomass ranged from 1.8 Mg·ha^–1 in the middle-aged stand to 3.5 Mg·ha^–1 in the young stand. Forest floor biomass increased steady with stand age, ranging from 14.9 to 23.0 Mg·ha^–1. The highest mean C concentration across the chronosequence was found in tree branch while the lowest mean C concentration was found in forest floor. The observed C stock of the aboveground tree, shrub, forest floor, and mineral soil increased with increasing stand age, whereas the herb C stock showed a decreasing trend with a sigmoid pattern. The C stock of forest ecosystem in young, middle-aged, immature, and mature stands were 178.1, 236.3, 297.7, and 359.8 Mg C ha^–1, respectively, greater than those under similar aged P. tabulaeformis forests in China. These results are likely to be integrated into further forest management plans and generalized in other contexts to evaluate C stocks at the regional scale.     

Landscape Patterns of Sapling Density,Leaf Area,and Aboveground Net Primary Production in Postfire Lodgepole Pine Forests,Yellowstone National Park (USA)

Causes and implications of spatial variability in postfire tree density and understory plant cover for patterns of aboveground net primary production (ANPP) and leaf area index (LAI) were examined in ninety 11-year-old lodgepole pine (Pinus contorta var. latifolia Engelm.) stands across the landscape of Yellowstone National Park (YNP), Wyoming, USA. Field studies and aerial photography were used to address three questions: (1) What is the range and spatial pattern of lodgepole pine sapling density across the burned Yellowstone landscape and what factors best explain this variability? (2) How do ANPP and LAI vary across the landscape and is their variation explained by abiotic factors, sapling density, or both? (3) What is the predicted spatial pattern of ANPP and LAI across the burned Yellowstone landscape? Stand density spanned six orders of magnitude, ranging from zero to 535,000 saplings ha^?1, and it decreased with increasing elevation and with increasing distance from unburned forest (r ²?=?0.37). Postfire densities mapped from 1:30,000 aerial photography revealed that 66% of the burned area had densities less than 5000 saplings ha^?1 and approximately 25% had densities greater than 10,000 saplings ha^?1; stand density varied spatially in a fine-grained mosaic. New allometric equations were developed to predict aboveground biomass, ANPP, and LAI of lodgepole pine saplings and the 25 most common herbaceous and shrub species in the burned forests. These allometrics were then used with field data on sapling size, sapling density, and percent cover of graminoid, forb, and shrub species to compute stand-level ANPP and LAI. Total ANPP averaged 2.8 Mg ha^?1y^?1 but ranged from 0.04 to 15.12 Mg ha^?1y^?1. Total LAI averaged 0.80 m² m^?2 and ranged from 0.01 to 6.87 m² m^?2. Variation in ANPP and LAI was explained by both sapling density and abiotic factors (elevation and soil class) (ANOVA, r ²?=?0.80); abiotic variables explained 51%–54% of this variation. The proportion of total ANPP contributed by herbaceous plants and shrubs declined sharply with increasing sapling density (r ²?=?0.72) and increased with elevation (r ²?=?0.36). However, total herbaceous productivity was always less than 2.7 Mg ha^?1 y^?1, and herbaceous productivity did not compensate for tree production when trees were sparse. When extrapolated to the landscape, 68% of the burned landscape was characterized by ANPP values less than 2.0 Mg ha^?1y^?1, 22% by values ranging from 2 to 4 Mg ha^?1y^?1, and the remaining 10% by values greater than 4 Mg ha^?1y^?1. The spatial patterns of ANPP and LAI were less heterogeneous than patterns of sapling density but still showed fine-grained variation in rates. For some ecosystem processes, postfire spatial heterogeneity within a successional stage may be similar in magnitude to the temporal variation observed through succession.     

Observations and modeling of aboveground tree carbon stocks and fluxes following a bark beetle outbreak in the western United States

Bark beetle epidemics result in tree mortality across millions of hectares in North America. However, few studies have quantified impacts on carbon (C) cycling. In this study, we quantified the immediate response and subsequent trajectories of stand‐level aboveground tree C stocks and fluxes using field measurements and modeling for a location in central Idaho, USA that experienced an outbreak of mountain pine beetle (Dendroctonus ponderosae Hopkins). We measured tree characteristics in lodgepole pine (Pinus contorta) plots spanning a range of structure and mortality conditions. We then initialized the forest vegetation simulator, an individual tree‐based model, with these measurements and simulated the response of aboveground production of C fluxes as well as trajectories of C stocks and fluxes in the coming decades. Mountain pine beetles killed up to 52% of the trees within plots, with more larger trees killed. C stocks in lodgepole pine were reduced by 31–83% following the outbreak, and plot‐level C fluxes decreased 28–73%. Modeled C stocks increased nearly continuously following the infestation, recovering to preoutbreak levels in 25 years or less. Simulated aboveground tree C fluxes increased following the immediate postoutbreak decrease, then subsequently declined. Substantial variability of C stocks and fluxes among plots resulted from the number and size of killed and surviving trees. Our study illustrates that bark beetle epidemics alter forest C cycling unlike stand‐replacement wildfires or clear‐cut harvests, due in part to incomplete mortality coupled with the preference by beetles for larger trees. The dependency of postoutbreak C stocks and fluxes on stand structure suggests that C budget models and studies in areas experiencing mountain pine beetle disturbances need to include size distribution of trees for the most accurate results.     

Stabilizing effects of diversity on aboveground wood production in forest ecosystems: linking patterns and processes

Both theory and evidence suggest that diversity stabilises productivity in herbaceous plant communities through a combination of overyielding, species asynchrony and favourable species interactions. However, whether these same processes also promote stability in forest ecosystems has never been tested. Using tree ring data from permanent forest plots across Europe, we show that aboveground wood production is inherently more stable through time in mixed‐species forests. Faster rates of wood production (i.e. overyielding), decreased year‐to‐year variation in productivity through asynchronous responses of species to climate, and greater temporal stability in the growth rates of individual tree species all contributed strongly to stabilising productivity in mixed stands. Together, these findings reveal the central role of diversity in stabilising productivity in forests, and bring us closer to understanding the processes which enable diverse forests to remain productive under a wide range of environmental conditions.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

A Decade of Streamwater Nitrogen and Forest Dynamics after a Mountain Pine Beetle Outbreak at the Fraser Experimental Forest,Colorado

Forests of western North America are currently experiencing extensive tree mortality from a variety of bark beetle species, and insect outbreaks are projected to increase under warmer, drier climates. Unlike the abrupt biogeochemical changes typical after wildfire and timber harvesting, the outcomes of insect outbreaks are poorly understood. The mountain pine bark beetle (Dendroctonus ponderosae) began to attack lodgepole pine (Pinus contorta) at the Fraser Experimental Forest in 2002 and spread throughout the research area by 2007. We compared streamwater nitrogen (N) from 2003 through 2012 with data from the previous two decades in four watersheds with distinct forest management histories, stand structures, and responses to the beetle outbreak. Watersheds dominated by old-growth had larger trees and lost 85% of overstory pine and 44% of total basal area to bark beetles. In contrast, managed watersheds containing a mixture of second-growth (30–60 year old) and old-growth (250- to 350-year old) had higher density of subcanopy trees, smaller mean tree diameter, and lower bark beetle-induced mortality (~26% of total basal area). Streamwater nitrate concentrations were significantly higher in old-growth watersheds during the outbreak than pre-outbreak levels during snowmelt and base flow seasons. In mixed-age stands, streamwater nitrate concentrations were unaffected by the outbreak. Beetle outbreak elevated inorganic N export 43 and 74% in two old-growth watersheds though the amounts of N released in streamwater were low (0.04 and 0.15 kg N ha^?1) relative to atmospheric inputs (<2% of annual N deposition). Increased height, diameter, and foliar N of measured in residual live trees augmented demand for N, far in excess of the change in N export during the outbreak. Reallocation of soil resources released after pine mortality to overstory and understory vegetation helps explain high nutrient retention in watersheds affected by bark beetle outbreaks.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

Soil inoculation of lodgepole pine seedlings alters root-associated fungal communities but does not improve seedling performance in beetle-killed pine stands

Extensive tree mortality in forests can change the community composition of soil fungi altering seedling establishment, a process critical to forest restoration. Disturbances that result in the loss of ectomycorrhizal fungi, in particular, may impede the establishment of tree species reliant on these symbionts for their survival. Inoculation of seedlings with soil from intact forests may improve the establishment of seedlings in such disturbances but the method has rarely been tested in the field. Here, we assess whether soil inoculation improves lodgepole pine (Pinus contorta var. latifolia) seedling performance in conspecific stands with high levels of tree mortality caused by a mountain pine beetle (Dendroctonus ponderosae) outbreak and whether underlying soil type modifies inoculation effects. We first inoculated seedlings in a growth chamber with small amounts of soils (5% volume) originating from either intact (<10%) or “beetle-killed” (>70% pine basal area killed) conspecific stands or added no soil inoculum and, after 4 months, transplanted them into 15 beetle-killed stands. After two growing seasons, root-associated fungal communities of seedlings receiving inoculum from intact stands differed in composition from those receiving inoculum from beetle-killed stands or no inoculum. However, inoculation had no effect on seedling survival, height, or biomass. Site properties, including soil texture and the resident fungal community composition, overwhelmed the effect of soil inoculation on seedling performance. Seedling survival and shoot mass was higher in sandy than loamy soils. Restoration to improve seedling performance in beetle-killed stands should consider stand-level treatments as soil inoculation at the level evaluated was ineffective.     

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

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

Resistance to wildfire and early regeneration in natural broadleaved forest and pine plantation

The response of an ecosystem to disturbance reflects its stability, which is determined by two components: resistance and resilience. We addressed both components in a study of early post-fire response of natural broadleaved forest (Quercus robur, Ilex aquifolium) and pine plantation (Pinus pinaster, Pinus sylvestris) to a wildfire that burned over 6000 ha in NW Portugal. Fire resistance was assessed from fire severity, tree mortality and sapling persistence. Understory fire resistance was similar between forests: fire severity at the surface level was moderate to low, and sapling persistence was low. At the canopy level, fire severity was generally low in broadleaved forest but heterogeneous in pine forest, and mean tree mortality was significantly higher in pine forest. Forest resilience was assessed by the comparison of the understory composition, species diversity and seedling abundance in unburned and burned plots in each forest type. Unburned broadleaved communities were dominated by perennial herbs (e.g., Arrhenatherum elatius) and woody species (e.g., Hedera hibernica, Erica arborea), all able to regenerate vegetatively. Unburned pine communities presented a higher abundance of shrubs, and most dominant species relied on post-fire seeding, with some species also being able to regenerate vegetatively (e.g., Ulex minor, Daboecia cantabrica). There were no differences in diversity measures in broadleaved forest, but burned communities in pine forest shared less species and were less rich and diverse than unburned communities. Seedling abundance was similar in burned and unburned plots in both forests. The slower reestablishment of understory pine communities is probably explained by the slower recovery rate of dominant species. These findings are ecologically relevant: the higher resistance and resilience of native broadleaved forest implies a higher stability in the maintenance of forest processes and the delivery of ecosystem services.     

连栽杉木林林下植被生物量动态格局

用空间一致时间连续的定位研究方法,在湖南会同杉木林生态系统国家野外科学观测研究站试验基地的第2集水区,对连栽杉木林林下植被生物量进行了12 a的监测,研究了林下植被种类的变化、生物量动态特征、生物量的组成与分布变化格局。结果表明:连栽杉木林在14a生长发育过程中,林下植物种类呈现波动性的减少趋势,其中木本植物物种数下降率为40.0%,草本植物物种数下降率为47.1%。林下植被生物量由杉木林3年生29.48 t/hm²下降至14年生的2.53 t/hm²,其中木本植物生物量由7.07 t/hm²,下降至1.25 t/hm²,下降了82.3%;草本植物由22.41 t/hm²,下降至1.28 t/hm²,下降了94.3%。在此期间,木本与草本植物生物量的高低均出现波动现象。3年生杉木林下木本植物以乔木树种生物量6068.97 kg/hm²最高,占总生物量85.88%,藤本植物生物量736.97 kg/hm²为次,占10.44%,灌木植物生物量259.87 kg/hm²最低,仅占3.68%。14年生杉木林下木本植物以灌木植物生物量881.87 kg/hm²为首,占总生物量70.73%,藤本植物生物量247.07 kg/hm²为次,占19.82%,乔木树种生物量117.87 kg/hm²最少,只占9.45%。3年生杉木林下草本植物以蕨类植物生物量8391.44 kg/hm²最高,占总生物量的37.44%,过路黄生物量36.77 kg/hm²最低,仅占0.16%。杉木14年生时,以芒生物量573.00 kg/hm²最大,占总生物量44.78%,金毛耳草生物量2.93 kg/hm²最小,仅占0.23%。研究结果,可为研究杉木林养分循环、碳平衡、维护和提高林地地力及可持续经营管理提供科学依据。     

Lodgepole pine forest age class dynamics and susceptibility to mountain pine beetle attack

Because mountain pine beetle attack mature pine stands, an understanding of forest age class dynamics is important to managing forests within the distribution of the beetle. The assumed theoretical negative exponential forest age distribution provides an estimate when ecosystem dynamics are in equilibrium. This study investigates the dynamics of forest age distribution for non-equilibrium ecosystem dynamics, which result primarily from large and irregular stand-replacement fire disturbances that alter the forest age distribution. A model experiment using the SEM-LAND model on a 1 million ha lodgepole pine forest landscape was conducted to estimate how the proportion of susceptible area could be influenced by different fire regimes. The results of the simulation suggest that the temporal dynamics of the area susceptible to mountain pine beetle attack are complex and depend on the fire history of the study area, if the area is experiencing large and irregular stand-replacement fires. The age range of the lodgepole pine forest stands susceptible to mountain pine beetle attack might significantly affect the estimate of the area susceptible to attack.     

南方红壤区3种典型森林恢复方式对植物群落多样性的影响

合理的森林恢复方式能提高植物多样性,进而提高生态系统服务功能.在我国南方红壤区研究了3种典型森林恢复方式(引进种恢复的湿地松(Pinus elliottii)人工林本地种恢复的马尾松(Pinus massoniana)人工林和自然恢复的天然次生林)的植物区系构成和植物群落多样性.结果表明:(1)湿地松人工林有155种植物,隶属66科118属,马尾松人工林有137种植物,隶属59科97属;天然次生林有226种植物,隶属86科160属,3种森林恢复方式的乔木层、灌木层和草本层的优势科属明显不同,马尾松人工林的优势物种和天然次生林更相似;(2)天然次生林的植物区系基本构成、植物区系类型种类高于马尾松人工林和湿地松人工林,并且天然次生林的温带成分比例高于湿地松人工林;(3)恢复方式对植物群落的多样性指数有显著影响,天然次生林的物种丰富度、辛普森指数明显高于马尾松人工林和湿地松人工林,两种人工林之间差异不显著;(4)3种森林恢复方式的植物群落结构存在显著差异,相比湿地松人工林,马尾松人工林的植物群落组成与天然次生林更相似.总之,自然恢复的天然次生林植物群落多样性高于人工恢复的马尾松人工林和湿地松人工林,本地种马尾松人工林在维持区域植物群落结构功能上优于引进种湿地松人工林.