Effects of Human Subsistence Activities on Forest Birds in Northern Kenya

Abstract: Indigenous tribes and conservation biologists may have common goals and may be able to collaborate on the maintenance of biodiversity, but few researchers have evaluated the impacts and potential benefits of human subsistence activities. I studied the effects of subsistence activities (primarily wood collection) of nomadic pastoralists in 3 Afromontane forests of northern Kenya. In surveys of 404, 25‐m‐radius plots, I recorded vegetation structure and composition of the forest bird community. Plots with higher levels of human activity had significantly different vegetation structure, with more‐open canopies, more grass, and fewer tree stems. Nectarivores (abundance +231%) and aerial insectivores (+66%) were more abundant in plots with more‐intense wood collecting than in plots with less human activity, whereas abundance of forest specialists (?28%) decreased in plots with more‐intense human activity. Abundance of 58% of the bird species either increased or decreased significantly in plots with more‐intense human activity. Generally, the number of individuals of forest specialists decreased (6 of 7 species showed significant responses) and the number of individuals of edge and nonforest species increased with increasing human activity. Canonical correspondence analysis showed that an intensification of human activities would favor nectarivores, aerial insectivores, granivores, and omnivores and would negatively affect large‐sized, ground‐foraging species and arboreal frugivores. Subsistence human activities favored the invasion of forest by edge species at the expense of forest specialists; thus, further intensification of forest exploitation by local peoples is not recommended. At the same time, however, subsistence activities in northern Kenya forests appeared to increase the structural diversity of the vegetation and provided suitable habitat for part (but not all) of the forest avifauna, which suggests that subsistence human activities may have a role in the maintenance of bird diversity.     

Comparative diversity and heavy metal biosorption of myxomycetes from forest patches on ultramafic and volcanic soils

Ultramafic and volcanic soils are exploited for industrial activities such as mining due to their high metal content, thus it is important that species in these areas are documented before irreversible environmental damage sets in. In this study, aerial and ground leaf litter, dead vines and twigs from six forest patches on volcanic and ultramafic soils in the provinces of Bataan, Pangasinan and Zambales in the Philippines were cultured in moist chambers (MC) and assessed for myxomycete diversity. From the 77% positive MC for myxomycetes, a total of 40 species from 14 genera were identified. Despite the higher heavy metal content, forest patches on ultramafic soils had greater species diversity as compared to volcanic soils. In this study, 10 species were abundant in both forest patches, namely Arcyria cinerea, Diachea leucopodia, Diderma effusum, D. hemisphaericum, Didymium ochroideum, Perichaena chrysosperma, P. corticalis, P. depressa, P. dictyonema and Physarum melleum. Selected myxomycetes tested for Cr and Mn content had equal or higher heavy metal levels than that of their leaf substrate. The study hypothesised that the presence of Mn⁷⁺ in fruiting bodies of myxomycetes was due to the phagocytosis of food bacteria inhabiting the substrates on the forest soil laden with heavy metal.     

Numerical study of wave–current–vegetation interaction in coastal waters

Research on interactions among wave, current, and vegetation has received increasing attention. An explicit depth-averaged hydrodynamic model coupled with a wave spectral model (CMS-wave) was proposed in this study in order to simulate the wave and wave-induced current in coastal waters. The hydrodynamic model was based on the finite volume method while the intercell flux was computed by employing the Harten–Lax–van Leer approximate Riemann solver to investigate the dry-to-wet interface, and the drag force of vegetation was modeled as the sink terms in the momentum equations. The CMS-wave model was used to investigate the non-breaking and the breaking random waves propagation in vegetation fields. Afterwards, an empirical wave energy dissipation term with plant effect was derived to represent the resistance induced by aquatic vegetation in the wave-action balance equation. The established model was calibrated and validated with both the experimental and field data. The results showed that the wave height decreased significantly along the wave propagation direction in the presence of vegetations. The sensitivity analysis for the plant density, the wave height, and the water depth were performed by comparing the numerical results for the wave height attenuation. In addition, wave and wave-induced current through a finite patch of vegetation in the surf zone were investigated as well. The strong radiation stress gradient could be produced due to the variation of the energy dissipation by vegetation effect in the nearshore zone, which impacted the direction and amplitude of the longshore current. The calculated results showed that the coupling model had good performance in predicting wave propagation and the current over vegetated water regions.     

Evaluating pollution-related damage and restoration success in urban forests with participatory monitoring and digital tools

Peri-urban forest monitoring requires indicators of vegetation damage. An example is the sacred fir (Abies religiosa) forests surrounding Mexico City, which have been heavily exposed to tropospheric ozone, a harmful pollutant, for over 4 decades. We developed a participatory monitoring system with which local community members and scientists generated data on ozone tree damage. Santa Rosa Xochiac rangers (13) used the digital tool KoboToolBox to record ozone damage to trees, tree height, tree ages, tree condition, tree position, and whether the tree had been planted. Thirty-five percent of the trees (n = 1765) had ozone damage. Younger trees had a lower percentage of foliage damaged by ozone than older trees (p < 0.0001), and asymptomatic trees tended to be younger (p < 0.0001). Symptomatic trees were taller than asymptomatic trees of the same age (R²_c = 0.43, R²_m = 0.27). Involving local communities facilitated forest monitoring and using digital technology improved data quality. This participatory system can be used to monitor forest condition change over time and thus aids restoration efforts driven by government or local communities’ interests, facilitating local decision-making.     

Simulation of wetlands forest vegetation dynamics

A computer program, SWAMP, was designed to simulate the effects of flood frequency and depth to water table on southern wetlands forest vegetation dynamics. By incorporating these hydrologic characteristics into the model, forest vegetation and vegetation dynamics can be simulated. The model, based on data from the White River National Wildlife Refuge near De Witt, Arkansas, “grows” individual trees on a 20 x 20-m plot taking into account effects on the tree growth of flooding, depth to water table, shade tolerance, overtopping and crowding, and probability of death and reproduction. A potential application of the model is illustrated with simulations of tree fruit production following flood-control implementation and lumbering.     

Standing crop and leaf area index of the sublittoral Laminaria species near Helgoland

Standing crop values of Laminaria species sampled between 0 to 2 m depth in the sublittoral of Helgoland, a small rocky island in the southern North Sea, are similar to those reported by other workers from Norwegian or Scottish coasts. Below 2 m water depth, however, standing crop values are found to be reduced to a far greater extent with increasing depth than in the Norwegian or Scottish areas studied. Maximum standing crop values are not found in Laminaria digitata and L. saccharina growing just below the low water mark of spring tides, but in the upper L. hyperborea forest at 2 m depth. Population density and leaf area index (LAI) decrease with increasing depth (but not the average weight of individuals constituting the L. hyperborea forest). It is suggested that the depth at which LAI becomes 1 represents an important ecological threshold, since growth rates of L. hyperborea specimens appear to decrease at greater depths but not to increase above the threshold. Self-shading might equalize, to a certain extent, the quantity of underwater irradiance available to an individual phylloid at all depths at which a close vegetation of Laminaria species occurs.     

Biodiversity Loss in Latin American Coffee Landscapes: Review of the Evidence on Ants,Birds, and Trees

Abstract: Studies have documented biodiversity losses due to intensification of coffee management (reduction in canopy richness and complexity). Nevertheless, questions remain regarding relative sensitivity of different taxa, habitat specialists, and functional groups, and whether implications for biodiversity conservation vary across regions. We quantitatively reviewed data from ant, bird, and tree biodiversity studies in coffee agroecosystems to address the following questions: Does species richness decline with intensification or with individual vegetation characteristics? Are there significant losses of species richness in coffee‐management systems compared with forests? Is species loss greater for forest species or for particular functional groups? and Are ants or birds more strongly affected by intensification? Across studies, ant and bird richness declined with management intensification and with changes in vegetation. Species richness of all ants and birds and of forest ant and bird species was lower in most coffee agroecosystems than in forests, but rustic coffee (grown under native forest canopies) had equal or greater ant and bird richness than nearby forests. Sun coffee (grown without canopy trees) sustained the highest species losses, and species loss of forest ant, bird, and tree species increased with management intensity. Losses of ant and bird species were similar, although losses of forest ants were more drastic in rustic coffee. Richness of migratory birds and of birds that forage across vegetation strata was less affected by intensification than richness of resident, canopy, and understory bird species. Rustic farms protected more species than other coffee systems, and loss of species depended greatly on habitat specialization and functional traits. We recommend that forest be protected, rustic coffee be promoted, and intensive coffee farms be restored by augmenting native tree density and richness and allowing growth of epiphytes. We also recommend that future research focus on potential trade‐offs between biodiversity conservation and farmer livelihoods stemming from coffee production.     

Short reviews by the Editor

Soil desiccation is a major issue limiting development and sustainability of forest vegetation in the Loess Plateau of China. Better understanding of the mechanisms of soil desiccation in the Loess Plateau can help scientists and forest managers improve vegetation management practices. The arid soil layer is the ecological aftermath of intense soil desiccation due to disturbed plant succession and soil water reduction. The formation and types of arid soil layer in the Loess Plateau were investigated to determine major causes of soil desiccation and its impact on forest vegetation. The negative effects of soil desiccation on the ecological environment and forest vegetation mainly include drying microclimate, degrading soil quality, poor vegetation growth, difficult forest renewal from natural seed banks, making it even more difficult to reforest forest lands and grasslands following plant senescence. Low precipitation, high evaporation, soil and water losses, improper selection of vegetation types, and too high population density of trees are probably the major reasons for the arid soil layer. Proper selection of vegetation types, adjusting tree density and other management practices can reduce the negative effects of the arid soil layer on forest vegetation.     

Elemental composition of Sal forest soils around Chhotanagpur Plateau,India

In recent decades, the Sal-dominated tropical deciduous forests (TDFs) were experiencing loss of regeneration due to lack of seed germination of native species. To understand this ecological problem, the present study dealt with the spatial and temporal variations in biogeochemical parameters of Sal forest soils. Twenty soil samples were collected from four TDF sites in the Chota Nagpur Plateau, India. The inventories of tree species in three life-cycle stages (seedling, sapling and adult) were carried out in four sites. Site-I (290?stems?ha^?1) showed occurrence of only one species (Shorea robusta) without seedling and sapling. The pH of forest soils was acidic to slightly acidic in nature. Seasonal variations in C/N ratio showed rapid mineralization of organic matter in the rainy season. ANOVA revealed that the degree of seasonal variation caused significant difference in pH, soil organic carbon, soil organic matter, total carbon, Al, Fe, Ca, Mg, Mn and P across four sites. The strong correlation of P with pH, Al, Fe, Mg and Mn indicated highly oxidised and exchangeable nature of dry forest soils. Factor analysis of soil parameters revealed that the different combinations of edaphic conditions in different seasons were important in the distribution of the dry tropical forests communities in Chota Nagpur Plateau.     

大兴安岭南部温带山杨天然次生林不同生长阶段生物量及碳储量

基于内蒙古赛罕乌拉森林生态系统定位研究站山杨（Populus davidiana Dode）天然次生林幼龄林、中龄林、近熟林、成熟林及过熟林生物量调查,探讨了不同龄组山杨天然次生林单株木、林分、林下植被和枯落物的生物量及群落碳储量的时空变化规律。结果表明：随林龄的增大,山杨天然次生林木和各器官生物量总体呈增加趋势,树干所占比例增加,中龄林增加尤为明显;林下植被层、枯落物层生物量随林龄增大呈增加趋势。群落总碳储量的空间分布序列是：乔木层〉枯落物层〉林下植被层。幼龄林、中龄林、近熟林、成熟林和过熟林群落的碳储量分别为27.146 6、53.545 1、60.889 8、77.915 8、79.135 3t.hm-2,乔木层碳储量分别为22.206 5、47.215 7、52.056 3、68.445 3、68.773 1 t.hm-2,枯落物层和林下植被层碳储量平均值分别为5.814 4、2.172 7 t.hm-2。乔木层、枯落物层和林下植被层碳储量占总量的平均率分别为86.05%、10.39%和3.57%。研究认为山杨天然次生林群落碳储量随林龄增加的变化规律明显,碳汇潜力巨大;中龄林为碳储量增长迅速期,且持续较长一段时间,是林分管理的关键阶段;自然稀疏有利于促进林木生长,林分碳储量并未随林分密度下降而减小。     

Influence of human-induced disturbance on benthic microbial metabolism in the Pichavaram mangroves,Vellar–Coleroon estuarine complex,India

Large areas of mangroves in India are heavily disturbed by cattle grazing, hypersalinity, and other human-induced impacts. In two disturbed Avicennia marina forests and two undisturbed A. marina and Rhizophora apiculata forests in the Pichavaram mangroves of the Vellar–Coleroon estuarine complex, southeast India, we measured the rates and pathways of microbial decomposition of soil organic matter to determine if human impact is altering biogeochemical activity within these stands. Rates of total carbon oxidation (T_COX) were higher in the undisturbed A. marina forest (mean 199 mol C m^–2 year^–1) than in the two impacted stands (43 and 79 mol C m^–2 year^–1); rates of total carbon oxidation in the R. apiculata forest averaged 75 mol C m^–2 year^–1. Sulphate reduction (range 21–319 mmol S m^–2 day^–1) was the major decomposition pathway (65–85% of T_COX), except at the most disturbed forest (30% of T_COX). Rates of sulphate reduction at all sites peaked in sub-surface soils to a depth of about 1 m, leading to little carbon burial (3–5% of total C input). There was some evidence of measurable iron and manganese reduction in association with tree roots. Rates of microbial activity were rapid in comparison with rates measured in other mangrove soils, reflecting high rates of phytoplankton production and organic matter retention in this lagoon. Human-induced disturbance creates a sharp zonation of dry, hypersaline soil overlying less saline, wetter soil, suppressing surface microbial and root growth. We conclude that this vertical alteration of soil characteristics and biogeochemistry shifts the cycling of nutrients between trees and microbes to a disequilibrium state, partly explaining why mangroves are stunted in these declining forests.Communicated by G. F. Humphrey, Sydney     

杉木观光木混交林和杉木纯林群落细根生产力、分布及养分归还

研究了福建三明27a生杉机光木混交林和杉木群落细根（d＜2mm）的生产力、分布、和养分归还。结果表明，混交林细根生物量、N、P养分现存量分别为5．381thm^2、48．085kghm^-2和4．174kghm^-2，分别比杉木纯林增加17．4％、27．2％和20．0％，混交林林细根的年净生产力达4．124thm^-2a^-1，比纯林高出16．9％，混交林杉木和观光木细根均在表层土壤富集，而在较深层土壤再会得分布具镶嵌性；与混交林杉林相比，纯林杉木土吉表层细根量较少，最大分布层次下移，混交林中观光木细根的周转速率咪1．16，杉木为0．96和0．95；而林下植被层细根周转速率（1．46－1．52）均同于相应的乔木层，混交林细根的年死亡量、N和P养分年归还量分别达2．119thm^-2、18．559kghm^-21．565kgkhm^-2，分别是纯林的1．21倍、1．23倍和1．14 倍，其中林下植被细根占有较为重要位置，对细根分布与土壤性质的相关分析表明，细根的垂直分布与土壤全N的相关性最强（0．87－0．89）。     

Examining Fragmentation and Loss of Primary Forest in the Southern Bahian Atlantic Forest of Brazil with Radar Imagery

Abstract: The Atlantic rainforest of southern Bahia is one of the last remnants of the lowland forest of eastern Brazil that once covered the entire coastal area from Rio Grande do Norte to Rio Grande do Sul ( lat 8°–28° S) and has been deforested to a small fraction of its original cover (1–12%). All recent vegetation surveys have been based on optical satellite data, which is hampered by cloud cover and by southern Bahia's intricate mix of forest patches with other tree crops, especially cocoa. We describe the application of radar remote-sensing data to distinguish forest patches from cocoa planted in the shade of natural-forest trees. Radar, unlike optical sensors, is not obstructed by cloud cover and can acquire information about forest structure by penetrating into the vegetation canopy. The vegetation map generated from radar data clearly separates forest patches based on the degree of structural disturbance such as the density of shaded trees, the openness of the canopy, and the density of the monodominant Erythyrina shaded trees. The structural classification based on the radar data, and shown on the map, can help researchers assess the degree of fragmentation of the original Atlantic coastal forest and delineate areas of less disturbance with higher potential for conservation of biodiversity. This information can then be applied to conservation planning, especially the design and monitoring of nature reserves and the modeling of biological corridors.     

Ten years of land cover change on the California coast detected using Landsat satellite image analysis: Part 2—San Mateo and Santa Cruz counties

Landsat satellite imagery was analyzed to generate a detailed record of 10 years of vegetation disturbance and regrowth for Pacific coastal areas of San Mateo and Santa Cruz Counties. The Landsat Ecosystem Disturbance Adaptive Processing System (LEDAPS) methodology, a transformation of Tasseled-Cap data space, was applied to detected changes in perennial coastal shrubland, woodland, and forest cover from 1999 to 2009. Results showed several principal points of interest, within which extensive contiguous areas of similar LEDAPS vegetation change (either disturbed or restored) were detected. Regrowth of evergreen shrub and tree cover was prevalent along the several long stretches of the coast highway (CA Route 1) between the cities of Half Moon Bay and Santa Cruz. A number of state parks areas showed measurable vegetation restoration as well. The most prominent loss of perennial coastal vegetation over decade was in the Pescadero Marsh area, where the continued presence of levees has historically reduced flood conveyance capacity into and through the marshlands. Based on these examples, the LEDAPS methodology was determined to be capable of fulfilling much of the need for continual, low-cost monitoring of emerging changes to coastal ecosystems.     

Crossing the frontier: vertical transit rates of deep diving cormorants reveal depth zone of neutral buoyancy

Knowing the depth zone of neutral buoyancy of divers is important because buoyancy can determine how animals manage their energy budget. In this study, we estimate the depth zone of neutral buoyancy of free-ranging cormorants for the first time, using time-depth recorders. We discovered that vertical ascent rates of 12 Crozet and 15 Kerguelen diving blue-eyed shags (respectively Phalacrocorax melanogenis and P. verrucosus) slowed down considerably at the 50–60 m depth zone. We suggest this was due to birds trying to reach the surface from that point upwards using reduced locomotor activity because the force of buoyancy becomes greater than the force of gravity at that depth. The results show a shift of this depth zone in relation to maximum targeted dive depth, suggesting cormorants may control buoyancy through respiratory air volume adjustment. Interestingly, 60 m is close to the maximum depth zone reached by these two species during dives lasting 4 min, their estimated behavioural aerobic dive limit. This suggests that the decision to swim deeper has a direct consequence on the energy budget, with time spent recovering at the surface (time thus lost to foraging) strongly increasing relative to the preceding time of submergence. Resources found in deeper waters must be of sufficient quantity or quality to justify crossing the frontier of physical neutral buoyancy.     

Chemical composition of rivers and streams in the forest and savanna zones of Nigeria

The aim of this study is to investigate the differences in the chemical conditions of lotic waterbodies in the two major ecosystems in Nigeria, the forest and savanna zones. The forest waters were slightly acidic (mean±SD pH = 6.72±0.58) while the savanna waters were slightly alkaline (pH = 7.11±0.33). The cationic order of dominance in the forest waters was Na⁺ > Ca²⁺ > Mg²⁺ > K⁺ in contrast to Ca²⁺ > Mg²⁺ > Na⁺ > K⁺ in savanna waters. The forest waters were chlorided (typical of coastal and/or marine waters) whereas the savanna waters were carbonated in nature, typical of the worldwide freshwater. Organic carbon was significantly higher in forest waters than in the savanna waters (p < 0.05) while nutrient compounds were significantly higher in savanna waters than in forest waters. The seasonal variation of the chemical parameters was generally more evident in savanna than in forest waters. The differences in water quality between the two major vegetation zones reflect the differences in the biogeochemical processes and nutrient cycling that characterise forest and savanna ecosystems.     

Modelling and mapping topographic variations in forest soils at high resolution: A case study

This article examines the utility of a digitally derived cartographic depth-to-water (DTW) index to model and map variations in drainage, vegetation and soil type and select soil properties within a forested area (40 ha) of the Swan Hills, Alberta, Canada. This index was derived from a LiDAR (Light Detection and Ranging) derived digital elevation model (DEM), with at least 1 ground return per m². The resulting DTW pattern was set to be zero along all DEM-derived flow channels, each with a 4 ha flow-initiation threshold. Soil type (luvisol, gleysol, mesisol), drainage type (very poor to excessive), vegetation type (hydric to xeric) and forest floor depth were determined along hillslope transects. These determinations conformed more closely to the DEM-derived log₁₀(DTW) variations (R² > 60%) than to the corresponding variations of the widely used topographic wetness index (TWI) (R² < 25%). Setting log₁₀(DTW) thresholds to represent the wet to moist to dry transitions between vegetation, drainage and soil type enabled a high-resolution mapping of these types across the study area. Also determined were soil moisture content, coarse fragment and soil particle composition (sand, silt, clay), pH, total C, N, S, P, Ca, Mg, K, Fe, Al, Mn, Zn, and available Ca, Mg, K, P and NH₄, by soil layer type and depth. Most of these variables were also more correlated with log₁₀(DTW) than with TWI, with and without soil layer depth as an additional regression variable. These variables are, therefore, subject to topographic controls to at least some extent, and can be modelled and mapped accordingly, as illustrated for soil moisture, forest floor depth and pH across the study area, from ridge tops to depressions. Further examinations revealed that the DEM-produced DTW and TWI patterns complemented one another, with DTW delineating soils in relation to local water-table influences, and with TWI delineating where the water would flow and accumulate.     

The role of archaeophytes and neophytes in the Dutch coastal dunes

The position of alien plant species in the Dutch coastal dune vegetation is evaluated considering 12 archaeophytes and 20 neophytes (including one moss), all of widespread occurrence in the coastal area of the Netherlands. Almost all archaeophytes have become part of natural vegetation types. Open scrub communities, in particular Hippophae rhamnoides-Sambucus nigra scrub at the leeside of the outer dunes, and calciphilous moss-dominated pioneer vegetation are relatively rich in archaeophytes. Among neophyte vascular species a contrast is apparent between herbaceous and woody species. The majority of herbaceous neophytes are characteristic of man-disturbed habitats and are only infrequently observed in natural vegetation types. However, woody species (as well as the moss Campylopus introflexus) have entered into natural vegetation on a large scale and behave in an invasive way.     

Site specific factors have an overriding impact on Baltic dune vegetation change under low to moderate N-deposition—a case study from Hiddensee island

At three coastal dune sites at the island of Hiddensee, north-east Germany, vegetation cover was mapped during 2002 and compared to vegetation surveys from the late 1980s and 1930s. Abiotic and biotic factors, which have been identified as being critical for coastal dunes in former studies such as disturbance, salt spray or nutrient availability, were measured. Grazing and land-use history were reviewed by literature and interviews. Tall graminoid communities, mainly Carex arenaria, are a common vegetation unit today. Development, distribution of these dominances and possible causes for its occurrence have not been analysed. Generally, older successional vegetation units increased and pioneer stages decreased from the 1930s until 2002. At the geologically youngest site, the southern dunes, grass encroachment by Carex arenaria was highest (ca. 50% cover in 2002), and age and density of trees lower than at the older, central dunes. Land-use changes such as decrease in grazing pressure, additional feeding of livestock, increase in coastal protection measures and subsequent decrease in shifting sands as well as varying availability of groundwater and amount of salt spray are relevant factors for vegetation changes in coastal dunes over the past 70 years. Site-specific land-use differences such as livestock density and land-use history have a stronger influence than atmospheric N-pollution on the vegetation composition of these acidic, coastal dunes under low to moderate N-deposition loads of 6–8 kg N ha⁻¹ yr⁻¹.     

Simulation of interannual responses of trembling aspen stands to climatic variation and insect defoliation in western Canada

A carbon-based model has been developed to simulate responses of trembling aspen (Populus tremuloides Michx.) stands to interannual climatic variation and insect defoliation. The model is designed for medium time scale (10–100 years) simulations and requires only daily maximum and minimum temperature and precipitation as meteorological inputs. The modelling approach is similar to FOREST-BGC but includes additional processes known to be important in deciduous forests. These include removal of leaf area during outbreaks of forest tent caterpillar (Malacosoma disstria Hbn.), phenological changes in leaf area index, storage and allocation of non-structural carbohydrate and the contribution of understorey vegetation to evapotranspiration. The model was used for simulations of growth and mortality of biomass carbon in two mature aspen forests located in the climatically dry transition zone between the boreal forest and prairie grassland regions of Saskatchewan, Canada. Model inputs of annual defoliation intensity were based on historic records of insect defoliation and the incidence of light-coloured tree rings in disks or cores collected from aspen at each of the two sites. At both sites, moderately good correlations (r²=0.47–0.54) were obtained between modelled interannual changes in stem carbon growth and observed interannual changes in stem basal area increment obtained from tree-ring analysis. Model outputs of stem biomass carbon were found to be highly sensitive to parameters describing seasonal leaf area duration, insect defoliation intensity, photosynthesis and root respiration and carbohydrate allocation to growth versus storage.