N biogeochemistry and cycling in two well-drained savannas: a comparison between the Orinoco Basin (Llanos,Venezuela) and Ivory Coast (western Africa)

When a savanna burns, a decline in the input of organic matter and nutrients to the soil occurs. However, the existence of recurrent fires is a natural condition and N depletion by fire is not incompatible with the existence of savannas per se. Consequently, savanna vegetations have evolved under fires, implying a near to steady-state N budget. In some Australian and African ecosystems, N fixation appears to be insufficient to replace losses inducing soil-N depletion, whereas in neotropical Llanos and western African savannas, the N-fixation and precipitation seem to be enough to maintain production despite fires. This review presents information about well-drained savannas, namely Trachypogon savannas in Orinoco Llanos and Andropogonae savannas in Ivory Coast. The sites present similarities in climate and fire regime differing in soil parent material. A report on N budgets is presented. The budget was positive, since losses seem to be balanced by inputs in rainfall and biological fixation. Uncertainties in flux measures indicate that more work on those aspects are required. Results indicate that the estimated amounts of dinitrogen fixation and deposition seem to be enough to maintain plant production. Data support the hypothesis that the vegetation in the natural savanna has evolved under fire constraint.     

Are savannas patch-dynamic systems? A landscape model

Savannas are ecosystems characterized by the coexistence of woody species (trees and bushes) and grasses. Given that savanna characteristics are mainly formed from competition, herbivory, fire, woodcutting, and patchy soil and precipitation characteristics, we propose a spatially explicit model to examine the effects of the above-mentioned parameters on savanna vegetation dynamics in space and time. Furthermore, we investigate the effects of the above-mentioned parameters on tree–bush–grass ratios, as well as the degrees of aggregation of tree–bush–grass biomass. We parameterized our model for an arid savanna with shallow soil depth as well as a mesic one with generally deeper and more variable soil depths. Our model was able to reproduce savanna vegetation characteristics for periods of time over 2000 years with daily updated time steps. According to our results, tree biomass was higher than bush biomass in the arid savanna but bush biomass exceeded tree and grass biomass in the simulated mesic savanna. Woody biomass increased in our simulations when the soil's porosity values were increased (mesic savanna), in combination with higher precipitation. Savanna vegetation varied from open savanna to woodland and back to open savanna again. Vegetation cycles varied over ∼300-year cycles in the arid and ∼220-year cycles in the mesic-simulated savanna. Autocorrelation values indicated that there are both temporal and spatial vegetation cycles. Our model indicated cycling savanna vegetation at the landscape scale, cycles in cells, and patchiness, i.e. patch dynamics.     

Tree cover in sub-Saharan Africa: rainfall and fire constrain forest and savanna as alternative stable states

Savannas are known as ecosystems with tree cover below climate-defined equilibrium values. However, a predictive framework for understanding constraints on tree cover is lacking. We present (a) a spatially extensive analysis of tree cover and fire distribution in sub-Saharan Africa, and (b) a model, based on empirical results, demonstrating that savanna and forest may be alternative stable states in parts of Africa, with implications for understanding savanna distributions. Tree cover does not increase continuously with rainfall, but rather is constrained to low (<50%, "savanna") or high tree cover (>75%, "forest"). Intermediate tree cover rarely occurs. Fire, which prevents trees from establishing, differentiates high and low tree cover, especially in areas with rainfall between 1000 mm and 2000 mm. Fire is less important at low rainfall (<1000 mm), where rainfall limits tree cover, and at high rainfall (>2000 mm), where fire is rare. This pattern suggests that complex interactions between climate and disturbance produce emergent alternative states in tree cover. The relationship between tree cover and fire was incorporated into a dynamic model including grass, savanna tree saplings, and savanna trees. Only recruitment from sapling to adult tree varied depending on the amount of grass in the system. Based on our empirical analysis and previous work, fires spread only at tree cover of 40% or less, producing a sigmoidal fire probability distribution as a function of grass cover and therefore a sigmoidal sapling to tree recruitment function. This model demonstrates that, given relatively conservative and empirically supported assumptions about the establishment of trees in savannas, alternative stable states for the same set of environmental conditions (i.e., model parameters) are possible via a fire feedback mechanism. Integrating alternative stable state dynamics into models of biome distributions could improve our ability to predict changes in biome distributions and in carbon storage under climate and global change scenarios.     

Elephants, fire, and frost can determine community structure and composition in Kalahari Woodlands.

Fire, elephants, and frost are important disturbance factors in many African savannas, but the relative magnitude of their effects on vegetation and their interactions have not been quantified. Understanding how disturbance shapes savanna structure and composition is critical for predicting changes in tree cover and for formulating management and conservation policy. A simulation model was used to investigate how the disturbance regime determines vegetation structure and composition in a mixed Kalahari sand woodland savanna in western Zimbabwe. The model consisted of submodels for tree growth, tree damage caused by disturbance, mortality, and recruitment that were parameterized from field data collected over a two-year period. The model predicts that, under the current disturbance regime, tree basal area in the study area will decline by two-thirds over the next two decades and become dominated by species unpalatable to elephants. Changes in the disturbance regime are predicted to greatly modify vegetation structure and community composition. Elephants are the primary drivers of woodland change in this community at present-day population densities, and their impacts are exacerbated by the effects of fire and frost. Frost, in particular, does not play an important role when acting independently but appears to be a key secondary factor in the presence of elephants and/or fire. Unlike fire and frost, which cannot suppress the woodland phase on their own in this ecosystem, elephants can independently drive the vegetation to the scrub phase. The results suggest that elephant and fire management may be critical for the persistence of certain woodland communities within dry-season elephant habitats in the eastern Kalahari, particularly those dominated by Brachystegia spiciformis and other palatable species.     

Grass feedbacks on fire stabilize savannas

Savannas commonly consist of a discontinuous cover of overstory trees and a groundcover of grasses. Savanna models have previously demonstrated that vegetation feedbacks on fire frequency can limit the density of overstory trees, thereby maintaining savannas. Positive feedbacks of either savanna trees alone or trees and grasses together on fire frequency have been shown to result in a stable savanna equilibrium. Grass feedbacks on fire frequency, in contrast, have resulted in stable equilibria in either a grassland or forest state, but not in a savanna. These results, however, were derived from a system of differential equations that assumes that fire occurrence is strictly deterministic and that vegetation losses due to fire are continuous in time. We develop an alternative formulation of the grass-fire feedback model that assumes that fires are discrete and occur stochastically in time to examine the influence of these assumptions on the predicted state of the system. We show that incorporating fire as a discrete event can produce a recurring temporal refuge in which both grass and trees co-occur in a stable, bounded savanna. In our model, tree abundance is limited without invoking demographic bottlenecks in the transition from fire-sensitive to fire-resistant life history stages. An increasing strength of grass feedback on fire results in regular, predictable fires, which suggests that the system can also be modeled using a set of difference equations. We implement this discrete system using modified Leslie/Gower difference equations and demonstrate the existence of a bounded savanna state in this model framework. Our results confirm the potential for grass feedbacks to result in stable savannas, and indicate the importance of modeling fire as a discrete event rather than as a loss rate that is continuous in time.     

Postfire stand structure in a semiarid savanna: cross-scale challenges estimating biomass.

Algorithms relating remotely sensed woody cover to biomass are often the basis for large-scale inventories of aboveground carbon stocks. However, these algorithms are commonly applied in a generic fashion without consideration of disturbances that might alter vegetation structure. We compared field and remote sensing estimates of woody biomass on savannas with contrasting disturbance (fire) histories and assessed potential errors in estimating woody biomass from cover without considering fire history. Field surveys quantified multilayer cover (MLC) of woody and succulent plants on sites experiencing wildfire in 1989 or 1994 and on nearby unburned (control) sites. Remote sensing estimates of the woody cover fraction (WCF) on burned and control sites were derived from contemporary (2005) dry-season Landsat Thematic Mapper imagery (during a period when herbaceous cover was senescent) using a probabilistic spectral mixture analysis model. Satellite WCF estimates were compared to field MLC assessments and related to aboveground biomass using allometry. Field-based MLC and remotely sensed WCFs both indicated that woody cover was comparable on control areas and areas burned 11-16 years ago. However, biomass was approximately twofold higher on control sites. Canopy cover was a strong predictor of woody biomass on burned and control areas, but fire history significantly altered the linear cover-biomass relationship on control plots to a curvilinear relationship on burned plots. Results suggest predictions of woody biomass from "generic" two-dimensional (2-D) cover algorithms may underestimate biomass in undisturbed stands and overestimate biomass in stands recovering from disturbance. Improving the accuracy of woody-biomass estimates from field and/or remotely sensed cover may therefore require disturbance-specific models or detection of vegetation height and transforming 2-D vegetation cover to 3-D vegetation volume.     

Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-down effects

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant "bottom-up" effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating "top-down" effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the "landscape of fear" in which African savanna herbivores exist.     

Woody encroachment decreases diversity across North American grasslands and savannas

Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation (r2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.     

提高碳汇潜力：量化树种和造林模式对碳储量的影响

全球气候变化背景下,造林再造林固定的碳可以抵消温室气体减限排量。通过造林再造林增加森林面积可以增加林业碳汇,在土地面积有限的情况下,提高造林质量——在有限的造林面积上固定更多的碳是十分必要的。树种和造林模式的选择是增加森林生态系统碳汇的重要管理决策。文章综述了树种和造林模式对生态系统的碳储量的影响。树种从生物量的积累,凋落物和土壤碳储存,以及木材密度、碳贮存量等几个方面探讨其对生态系统碳库的影响。混交林能充分利用立地条件、改善树木营养状况,并且可以减少病虫害和森林火灾。同时分析了我国在森林经营方面存在的问题和改善途径,以期为该领域的研究提供参考。     

Small-scale fuel variation alters fire intensity and shrub abundance in a pine savanna

Small-scale variation in fire intensity and effects may be an important source of environmental heterogeneity in frequently burned plant communities. We hypothesized that variation in fire intensity resulting from local differences in fuel loads produces heterogeneity in pine savanna ground cover by altering shrub abundance. To test this hypothesis, we experimentally manipulated prefire fuel loads to mimic naturally occurring fuel-load heterogeneity associated with branch falls, needle fall near large pines, and animal disturbances in a frequently burned longleaf pine (Pinus palustris) savanna in Louisiana, USA. We applied one of four fuel treatments (unaltered control, fine-fuel removal, fine-fuel addition, wood addition) to each of 540 (1-m2) quadrats prior to growing-season prescribed fires in each of two years (1999 and 2001). In both years fuel addition increased (and fuel removal decreased) fuel consumption and maximum fire temperatures relative to unaltered controls. Fuel addition, particularly wood, increased damage to shrubs, increased shrub mortality, and decreased resprout density relative to controls. We propose that local variation in fire intensity may contribute to maintenance of high species diversity in pine savannas by reducing shrub abundance and creating openings in an otherwise continuous ground cover.     

Longleaf pine (Pinus palustris) and hardwood dynamics in a fire-maintained ecosystem: A simulation approach

Longleaf pine (Pinus palustris) savannas of the southeastern U.S. represent an archetype of a fire dependent ecosystem. They are known to have very short fire return intervals (∼1-3 years) that perpetuate understory plant diversity (up to 50 species m⁻²), support pine recruitment, and suppress fire sensitive hardwoods. Understanding the relationships that regulate longleaf and southern hardwoods is especially critical. With decreased fire frequency, insufficient intensity, or lack of underground competition, a woody mid-story rapidly develops, dominated by fire sensitive trees and shrubs that in-turn suppress more fire dependent species (including pine seedlings). This may occur in forest gaps, where pine-needle abundance is diminished, reducing fire spread potential. The interactions between longleaf pine, hardwoods, forest fuels, and fire frequency are complex and difficult to understand spatially. The objective of this study was to develop a spatially explicit longleaf pine-hardwood stochastic simulation model (LLM), incorporating tree demography, plant competition, and fuel and fire characteristics. Data from two longleaf pine study sites were used to develop and evaluate the model with the goal to incorporate simple site-specific calibration parameters for model versatility. Specific model components included pine seed masting, hardwood clonal sprouting, response to fire (re-sprouting, mortality), and tree density driven competition effects. LLM spatial outputs were consistent with observed forest gap dynamics associated with pine seedling establishment and hardwood encroachment. Changes in fire frequency (i.e., fire probability = 0.35-0.05) illustrated a shift in community structure from longleaf pine dominated to a hardwood dominated community. This approach to assessing model response may be useful in characterizing longleaf ecosystem resilience, especially at intermediate fire frequencies (e.g., 0.15) where the community may be sensitive to small changes in the fire regime. Height distributions and population densities were similar to in situ findings (field and LIDAR data) for both study sites. Height distributions output by the LLM illustrated fluctuations in population structure. The LLM was especially useful in determining knowledge gaps associated with fuel and fire heterogeneity, plant-plant interactions, population structure and its temporal fluctuations, and hardwood demography. This is the first known modeling work to simulate interactions between longleaf pine and hardwoods and provides a foundation for further studies on fire and forest management, especially in relation to ecological forestry practices, restoration, and site-specific applications.     

不同密度侧柏人工林碳储量变化及其机理初探

以徐州侧柏Platycladus orientalis（Linn）Franco人工林为研究对象,运用生物量转化方程及土壤调查数据探讨了1 679、2 250和3 074株.hm-2的3种密度对生态系统碳储量的影响及其机理。结果表明,①乔木层、土壤层和生态系统的碳储量均随林分密度的增加而明显减少,灌草层碳储量在低林分密度最大,而枯落物层碳储量在中林分密度最大。低林分密度生态系统的碳储量是94.11 t.hm-2,分别是中密度和高密度生态系统的碳储量1.19倍和1.28倍,而这种差异主要是由乔木层和土壤层碳储量差异引起的。②林分密度对细根生物量的影响不显著（P〉0.05）,而细根形态随林分密度的增加表现为低级根中1、2级根直径变粗,根长先变长后变短,比根长变短（P〈0.05）;而高级根中的5级根直径显著变细,根长和比根长变长（P〉0.05）。③林分密度对细根生物量的影响与乔木层、土壤层和生态系统碳储量的变化规律具有较高的一致性,均为低密度下最大,高密度下最小。因此,细根生物量可能是导致系统碳储量变化的主要因素之一。     

Predicting a ‘tree change’ in Australia's tropical savannas: Combining different types of models to understand complex ecosystem behaviour

In this study, key ecological modelling limitations of a process-based simulation model and a Bayesian network were reduced by combining the two approaches. We demonstrate the combined modelling approach with a case study investigating increases in woody vegetation density in northern Australia's tropical savannas. We found that by utilising the strengths of a simulation model and a Bayesian network we could both forecast future change in woody vegetation density and diagnose the reasons for current vegetation states. The local conditions of climate, soil characteristics and the starting population of trees were found to be more important in explaining the likelihood of change in woody vegetation density compared to management practices such as grazing pressure and fire regimes. We conclude that combining the strengths of a process and BN model allowed us to produce a simple model that utilised the ability of the process model to simulate ecosystem processes in detail and over long time periods, and the ability of the BN to capture uncertainty in ecosystem response and to conduct scenario, sensitivity and diagnostic analysis. The overall result was a model that has the potential to provide land managers with a better understanding of the behaviour of a complex ecosystem than simply utilising either modelling approach in isolation.     

Forest floor depth mediates understory vigor in xeric Pinus palustris ecosystems.

Longleaf pine (Pinus palustris) woodlands and savannas are among the most frequently burned ecosystems in the world with fire return intervals of 1-10 years. This fire regime has maintained high levels of biodiversity in terms of both species richness and endemism. Land use changes have reduced the area of this ecosystem by >95%, and inadequate fire frequencies threaten many of the remnants today. In the absence of frequent fire, rapid colonization of hardwoods and shrubs occurs, and a broad-leaved midstory develops. This midstory encroachment has been the focus of much research and management concern, largely based on the assumption that the midstory reduces understory plant diversity through direction competition via light interception. The general application of this mechanism of degradation is questionable, however, because midstory density, leaf area, and hardwood species composition vary substantially along a soil moisture gradient from mesic to extremely xeric sites. Reanalysis of recently reported data from xeric longleaf pine communities suggests that the development of the forest floor, a less conspicuous change in forest structure, might cause a decline in plant biodiversity when forests remain unburned. We report here a test of the interactions among fire, litter accumulation, forest floor development, and midstory canopy density on understory plant diversity. Structural equation modeling showed that within xeric sites, forest floor development was the primary factor explaining decreased biodiversity. The only effects of midstory development on biodiversity were those mediated through forest floor development. Boundary line analysis of functional guilds of understory plants showed sensitivity to even minor development of the forest floor in the absence of fire. These results challenge the prevailing management paradigm and suggest that within xeric longleaf pine communities, the primary focus of managed fire regime should be directed toward the restoration of forest floor characteristics rather than the introduction of high-intensity fires used to regulate midstory structure.     

Carbon sequestration of black locust forests in the Yellow River Delta region,China

The Yellow River Delta region in China is a land area of 1,200,000 ha with rich natural resources. Adverse environmental conditions, such as low rainfall and high salinity, promote the dominance of black locust trees for afforestation. With the increase of CO₂ in the atmosphere, this forest and others throughout the world have become valued for their ability to sequester and store carbon. Forests store carbon in aboveground biomass (i.e. trees), belowground biomass (i.e. roots), soils and standing litter crop (i.e. forest floor and coarse woody debris). There are well-developed methods to sample forest ecosystems, including tree inventories that are used to quantify carbon in aboveground tree biomass. Such inventories are used to estimate the types of roundwood products removed from the forest during harvesting. Based on standard plot inventories and stem analyses, carbon sequestration estimates of trees were 222.41 t ha^?1 for the Yellow River Delta region accounted for 67.12% of the whole forest. Similarly, carbon storage by herbaceous matter and soil was 0.50 and 50.34 t ha^?1, respectively. The results suggest that carbon sequestration in the forest ecosystem was performed by most of the forest, which plays an increasingly important role in sequestering carbon as the stand grows.     

Hyperspectral and LiDAR remote sensing of fire fuels in Hawaii Volcanoes National Park.

Alien invasive grasses threaten to transform Hawaiian ecosystems through the alteration of ecosystem dynamics, especially the creation or intensification of a fire cycle. Across sub-montane ecosystems of Hawaii Volcanoes National Park on Hawaii Island, we quantified fine fuels and fire spread potential of invasive grasses using a combination of airborne hyperspectral and light detection and ranging (LiDAR) measurements. Across a gradient from forest to savanna to shrubland, automated mixture analysis of hyperspectral data provided spatially explicit fractional cover estimates of photosynthetic vegetation, non-photosynthetic vegetation, and bare substrate and shade. Small-footprint LiDAR provided measurements of vegetation height along this gradient of ecosystems. Through the fusion of hyperspectral and LiDAR data, a new fire fuel index (FFI) was developed to model the three-dimensional volume of grass fuels. Regionally, savanna ecosystems had the highest volumes of fire fuels, averaging 20% across the ecosystem and frequently filling all of the three-dimensional space represented by each image pixel. The forest and shrubland ecosystems had lower FFI values, averaging 4.4% and 8.4%, respectively. The results indicate that the fusion of hyperspectral and LiDAR remote sensing can provide unique information on the three-dimensional properties of ecosystems, their flammability, and the potential for fire spread.     

Defining Historical Baselines for Conservation: Ecological Changes Since European Settlement on Vancouver Island,Canada

Abstract: Conservation and restoration goals are often defined by historical baseline conditions that occurred prior to a particular period of human disturbance, such as European settlement in North America. Nevertheless, if ecosystems were heavily influenced by native peoples prior to European settlement, conservation efforts may require active management rather than simple removal of or reductions in recent forms of disturbance. We used pre‐European settlement land survey records (1859–1874) and contemporary vegetation surveys to assess changes over the past 150 years in tree species and habitat composition, forest density, and tree size structure on southern Vancouver Island and Saltspring Island, British Columbia, Canada. Several lines of evidence support the hypothesis that frequent historical burning by native peoples, and subsequent fire suppression, have played dominant roles in shaping this landscape. First, the relative frequency of fire‐sensitive species (e.g., cedar [Thuja plicata]) has increased, whereas fire‐tolerant species (e.g., Douglas‐fir [Pseudotsuga menziesii]) have decreased. Tree density has increased 2‐fold, and the proportion of the landscape in forest has greatly increased at the expense of open habitats (plains, savannas), which today contain most of the region's threatened species. Finally, the frequency distribution of tree size has shifted from unimodal to monotonically decreasing, which suggests removal of an important barrier to tree recruitment. In addition, although most of the open habitats are associated with Garry oak (Quercus garryana) at present, most of the open habitats prior to European settlement were associated with Douglas‐fir, which suggests that the current focus on Garry oak as a flagship for the many rare species in savannas may be misguided. Overall, our results indicate that the maintenance and restoration of open habitats will require active management and that historical records can provide critical guidance to such efforts.     

Restoration of ecosystems for biodiversity and carbon sequestration: Simulating growth dynamics of brigalow vegetation communities in Australia

Restoration of abandoned and degraded ecosystems through enhanced management of mature remnant patches and naturally regenerating (regrowth) forests is currently being used in the recovery of ecosystems for biodiversity protection and carbon sequestration. Knowledge of long-term dynamics of these ecosystems is often very limited. Vegetation models that examine long-term forest growth and succession of uneven aged, mixed-species forest ecosystems are integral to the planning and assessment of the recovery process of biodiversity values and biomass accumulation. This paper examined the use of the Ecosystem Dynamics Simulator (EDS) in projecting growth dynamics of mature remnant brigalow forest communities and recovery process of regrowth brigalow thickets. We used data from 188 long-term monitored plots of remnant and regrowth forests measured between 1963 and 2010. In this study the model was parameterised for 34 tree and shrub species and tested with independent long-term measurements. The model closely approximated actual development trajectories of mature forests and regrowth thickets but some inaccuracies in estimating regeneration through asexual reproduction and mortality were noted as reflected in stem density projections of remnant plots that had a mean of absolute relative bias of 46.2 (±12.4)%. Changes in species composition in remnant forests were projected with a 10% error. Basal area values observed in all remnant plots ranged from 6 to 29 m² ha⁻¹ and EDS projections between 1966 and 2005 (39 years) were 68.2 (±10.9)% of the observed basal area. Projected live aboveground biomass of remnant plots had a mean of 93.5 (±5.9) t ha⁻¹ compared to a mean of 91.3 (±8.0) t ha⁻¹ observed in the plots. In regrowth thicket, the model produced satisfactory projections of tree density (91%), basal area (89%), height (87%) and aboveground biomass (84%) compared to the observed attributes. Basal area and biomass accumulation in 45-year-old regrowth plots was approximately similar to that in remnant forests but recovery of woody understorey was very slow. The model projected that it would take 95 years for the regrowth to thin down to similar densities observed in original or remnant brigalow forests. These results indicated that EDS can produce relatively accurate projections of growth dynamics of brigalow regrowth forests necessary for informing restoration planning and projecting biomass accumulation.     

几种不同更新的森林群落碳储量结构特征分析

森林更新是维持和扩大森林资源的主要途径,也是森林结构调整、森林可持续经营和构建多功能高效的森林生态系统的过程。在安徽南部的岭南林场,选择了马尾松（Pinus massoniana Lamb）人工林（MP）、杉木（Cunninghamia lanceolata）人工林（CF）、阔叶混交天然次生林（MB）和针阔混交人工次生林（MN）等4种具有典型代表性的森林群落类型,研究了不同更新方式形成的森林群落的碳储量结构特征。结果表明：（1）针阔混交次生林树干生物量密度最大,为（67.32±56.57）mg.hm-2,杉木人工林生物量密度最小,为（43.79±9.13）mg.hm-2,而马尾松树干生物量所占比例最大,为（64.04±1.49）%。阔叶混交次生林碳储量最高,为（126.47±90.75）mg.hm-2;（2）4种群落类型中,阔叶混交林与马尾松群落碳密度最大,分别为95.67和98.21mg.hm-2,杉木群落碳密度最小,为55.41 mg.hm-2。阔叶混交林中的灌木层生物量碳密度最大,为（17.438±24.627）mg hm-2,马尾松林的草本层和枯落层生物量碳密度最高,分别为（1.326±0.431）、（5.517±2.846）mg.hm-2;（3）阔叶混交林群落的地下碳储量最高,为（10.5±9.8）mg.hm-2,群落地下碳储量从大到小的顺序是阔叶混交林〉针阔混交林〉杉木林〉马尾松林。相应的群落地上碳储量从大到小的顺序是阔叶混交林〉针阔混交林〉马尾松林〉杉木林。杉木林根茎比（R/S）最大,为0.21±0.01,杉木林群落中的灌木层根茎比（R/S）最大,为1.61±0.11;（4）在阔叶混交林中,株数密度与乔木层、草本层的碳比例正相关。在杉木林群落中,平均胸径、株数密度与乔木层碳所占比例成负相关。除杉木林群落外,灌木层碳含量之比与胸径及密度等调查因子都呈负相关。     

Cryptic herbivores mediate the strength and form of ungulate impacts on a long-lived savanna tree

Plant populations are regulated by a diverse array of herbivores that impose demographic filters throughout their life cycle. Few studies, however, simultaneously quantify the impacts of multiple herbivore guilds on the lifetime performance or population growth rate of plants. In African savannas, large ungulates (such as elephants) are widely regarded as important drivers of woody plant population dynamics, while the potential impacts of smaller, more cryptic herbivores (such as rodents) have largely been ignored. We combined a large-scale ungulate exclusion experiment with a five-year manipulation of rodent densities to quantify the impacts of three herbivore guilds (wild ungulates, domestic cattle, and rodents) on all life stages of a widespread savanna tree. We utilized demographic modeling to reveal the overall role of each guild in regulating tree population dynamics, and to elucidate the importance of different demographic hurdles in driving population growth under contrasting consumer communities. We found that wild ungulates dramatically reduced population growth, shifting the population trajectory from increase to decline, but that the mechanisms driving these effects were strongly mediated by rodents. The impact of wild ungulates on population growth was predominantly driven by their negative effect on tree reproduction when rodents were excluded, and on adult tree survival when rodents were present. By limiting seedling survival, rodents also reduced population growth; however, this effect was strongly dampened where wild ungulates were present. We suggest that these complex interactions between disparate consumer guilds can have important consequences for the population demography of long-lived species, and that the effects of a single consumer group are often likely to vary dramatically depending on the larger community in which interactions are embedded.