Diversity and structure of the tree community of a fragment of tropical secondary forest of the brazilian Atlantic Forest domain 15 and 40 years after logging

Two adjacent tracts of tropical secondary forest, situated in Itambé do Mato Dentro, south-eastern Brazil, which had been regenerating for 15 and 40 years after clearing, were compared with the purpose of detecting differences in species diversity and composition, species guild composition (regeneration, stratification and dispersion), and stand structure. Four and three 1,125 m² plots laid on the 15- and 40-year-old stands, respectively, sampled 2,430 trees with diameter at the base of the stem > 5 cm. The number of species (S = 199) was high for this forest type and significantly higher for the older stand. Tree density was significantly higher in the younger stand, particularly for smaller trees, whereas the two stands did not differ in both basal area and volume per hectare. Trees of shade-tolerant and understory species were significantly more abundant in the older stand. Though sharing a large proportion of species (49%), the two stands differed significantly in the abundance of many species. Live stumps probably contributed to the relatively quick restoration of some forest characteristics, particularly species diversity, basal area and volume.

Alley cropping of willows and grassland for bioenergy provision: productivity, tree-crop interactions and energy balance

The demand for biomass for bioenergy has increased rapidly in industrialized countries in the recent years. Biogenic energy carriers are known to reduce CO2 emissions. However, the resource-inefficient production of biomass often caused negative impacts on the environment, e.g. biodiversity losses, nitrate leaching, and erosion. The detrimental effects evolved mainly from annual crops. Therefore, the aim of modern bioenergy cropping systems is to combine yield stability and environmental benefits by the establishment of mixed-cropping systems. A particular emphasis is on perennial crops which are perceived as environmentally superior to annual crops. Agroforestry systems represent such mixed perennial cropping systems and consist of a mix of trees and arable crops or grassland within the same area of land. Agroforestry practices vary across the globe and alley cropping is a type of agroforestry system which is well adapted to the temperate zone, with a high degree of mechanization. Trees are planted in rows and crops are planted in the alleyways, which facilitates their management by machinery. This study was conducted to examine a young alley cropping system of willows and two grassland mixtures for bioenergy provision under temperate climate conditions. The first part of the thesis identified possible competition effects between willows and the two grassland mixtures. Since light seemed to be the factor most affecting the yield performance of the understory in temperate agroforestry systems, a biennial in situ artificial shade experiment was established over a separate clover-grass stand to quantify the effects of shade. Data to possible below- and aboveground interactions among willows and the two grassland mixtures and their effects on productivity, sward composition, and quality were monitored along a tree-grassland interface within the alleys. In the second part, productivity of the alley cropping system was examined on a triennial time frame and compared to separate grassland and willow stands as controls. Three different conversion technologies (combustion of hay, integrated generation of solid fuel and biogas from biomass, whole crop digestion) were applied to grassland biomass as feedstock and analyzed for its energetic potential. The energetic potential of willow wood chips was calculated by applying combustion as conversion technique. Net energy balances of separate grassland stands, agroforestry and pure willow stands evaluated their energy efficiency. Results of the biennial artificial shade experiment showed that severe shade (80 % light reduction) halved grassland productivity on average compared to a non-shaded control. White clover as heliophilous plant responded sensitively to limited radiation and its dry matter contribution in the sward decreased with increasing shade, whereas non-leguminous forbs (mainly segetal species) benefited. Changes in nutritive quality could not be confirmed by this experiment. Through the study on interactions within the alleys of the young agroforestry system it was possible to outline changes of incident light, soil temperature and sward composition of clover-grass along the tree-grassland interface. Nearly no effects of trees on precipitation, soil moisture and understory productivity occurred along the interface during the biennial experiment. Considering the results of the productivity and the net energy yield alley cropping system had lower than pure grassland stands, irrespective of the grassland seed mixture or fertilization, but was higher than that for pure willow stands. The comparison of three different energetic conversion techniques for the grassland biomass showed highest net energy yields for hay combustion, whereas the integrated generation of solid fuel and biogas from biomass (IFBB) and whole crop digestion performed similarly. However, due to the low fuel quality of hay, its direct combustion cannot be recommended as a viable conversion technique, whereas IFBB fuels were of a similar quality to wood chip from willow.

Farm level tree planting in Pakistan: The role of farmers' perceptions and attitudes

The low proportion of forested land and continuing degradation of existing forest cover are serious threats to the sustainability of forestry in Pakistan. Farm forestry has been identified as a feasible solution, particularly in the plain areas. Applying the Theory of Planned Behaviour in a survey of 124 farmers in Dera Ismail Khan district of Pakistan's North West Frontier Province showed that farmers' willingness to grow trees on their farms is a function of their attitudes towards the advantages and disadvantages of growing trees, their perception of the opinions of salient referents and factors that encourage and discourage farm level tree planting. Farmers viewed farm forestry as economically beneficial and environmentally friendly. Tree planting was perceived as increasing income, providing wood for fuel and furniture, controlling erosion and pollution and providing shade for humans and animals. Farmers saw hindrance in agricultural operations and the harbouring of insects, pests and diseases as negative impacts of tree planting; however, these were outweighed by their perceptions of positive impacts. Tree growing decisions of farmers were influenced by the opinions of family members, owners/tenants, fellow farmers and village elders. The factors that significantly predicted farm level tree planting were availability of barren land, lack of markets, lack of nurseries and damage caused by animals and humans. Farm forestry programmes are more likely to be successful if they acknowledge and address the factors which underlie farmers' reasons for planting or not planting trees.

Variations in tree community composition and structure in a fragment of tropical semideciduous forest in southeastern Brazil related to different human disturbance histories

In an area of tropical seasonal semideciduous forest, the soil characteristics, floristic composition, physiognomic structure, and the distribution of three regeneration and three dispersal guilds were studied for four stands within the forest that had documented histories of varying degrees of human disturbance. The aim was to study forest regeneration in areas of preserved forest and secondary forest, with parts of both types of forest experiencing either 'intensive' or 'occasional' cattle trampling. The study was carried out in the Sebastiao Aleixo da Silva Ecological Station, Bauru, São Paulo State, Brazil. Two stands were called 'secondary' because they corresponded to forest tracts that were felled and occupied by crops and pastures in the past and then abandoned to forest regeneration ca. 40 years before this study. The other two stands, called 'preserved', corresponded to areas of the fragment where the forest has been maintained with only minor human impacts. The arboreal component of the tree community (diameter at breast height or dbh greater than or equal to 5 cm) was sampled in 20 plots of 40 m x 40 m, and the subarboreal component (diameter at the base of the stem or dbs < 5 cm and height greater than or equal to 0.5 m) in subplots of 40 m x 2 m. Physiognomic features, such as canopy height and density of climbing plants, were registered all over a 5 m x 5 m gridline laid on the sample plots. Soil bulk samples were collected for chemical and textural analyses. Most detected differences contrasted the secondary to the preserved forest stands. The soils of the secondary stands showed higher proportions of sand and lower levels of mineral nutrients and organic matter than those of the preserved stands, probably due to higher losses by leaching and erosion. Compared to the secondary stands, the preserved ones had higher proportions of tall trees, higher mean canopy height, lower species diversity, higher abundance of autochorous and shade-tolerant climax species, and lower abundance of pioneer and light-demanding climax species. Despite the high proportion of species shared by the preserved and secondary stands (108 out of 139), they differed consistently in terms of density of the most abundant species. on the other hand, the secondary and preserved stands held similar values for tree density and basal area, suggesting that 40 years were enough to restore these features. Effects of cattle trampling on the vegetation were detected for the frequency of trees of anemochorous and zoochorous species, which were higher in the stands under occasional and intensive cattle trampling, respectively. The density of thin climbers was lower in the stands with intensive trampling. (C) 2004 Elsevier B.V. All rights reserved.

Leaf Morphological Plasticity of Tree Species from Two Developmental Stages in Araucaria Forest

This study compared the morphological and anatomical variations of the leaves of four shade-tolerant tree species Allophylus edulis (St.-Hil.) Radlk (Sapindaceae), Casearia sylvestris Sw. (Salicaceae), Cupania vernalis Cambess. (Sapindaceae) and Luehea divaricata Mart. (Malvaceae) from a fragment of Araucaria forest in two developmental stages. Morphological and anatomical traits, such as leaf and tissue thickness, leaf area, leaf dry mass, specific leaf area, leaf density and stomata density were measured from 30 leaves of each developmental stage. The phenotypic plasticity index was also calculated for each quantitative trait. The results showed that the four species presented higher mean values ​​for specific leaf area and spongy/palisade parenchyma ratio at young stage, and higher mean values ​​for stomata density, total and palisade parenchyma thickness in the adult stage. The plasticity index demonstrated that L. divricata presented highest plasticity for both the morphological and anatomical traits while A. edulis displayed the lowest plasticity index. The results of this study indicated that the leaves of these species exhibited distinct morphological traits at each stage of development to cope with acting environmental factors.

Initial growth of bauhinia variegata trees under different colored shade nets and light conditions

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Stock of biomass and carbon in the montane mixes shade forest, Parana

The aim of this study was to estimate the stock of biomass and organic carbon in a montane mixed shade forest located near General Carneiro, PR. 20 plots of 12 m x 12 m were installed, in which all trees with a CBH (Circumference at Breast Height) >= 31.4 cm were felled. From these the following information was obtained: total height, commercial height (agreed as being the morphological inversion point in the natural forest and the height of the first live branch), CBH, identification and collection of herbarium specimens. For the quantification of biomass in the understory and roots, three subunits 1 m x 1 m in each sampling unit were installed (12 m x 12 m) arranged in the lower left corner, center and diagonal upper right corner. To quantify accumulated litter at random, eight samples in each sampling unit were collected (12 m x 12 m), using a metal device measuring 0.25 m x 0.25 m. The montane mixed shade forest has more than 85% of its total biomass and total organic carbon stored in above ground plant structures. The total stock of organic carbon found in this study (104.7 Mg ha(-1)) demonstrates the importance of maintaining and preserving natural ecosystems as a way of maintaining this stock of organic carbon fixed in plant biomass.

Recruitment dynamics of the grove-dominant tree Microberlinia bisulcata in African rain forest: extending the light response versus adult longevity trade-off concept

In groves of ectomycorrhizal caesalpiniaceous species in the Atlantic coastal forest of Central Africa the dominant tree Microberlinia bisulcata, which is shade-intolerant as a seedling but highly light-responding as a sapling, shows very limited regeneration. M. bisulcata saplings were mapped in an 82.5-ha plot at Korup and found to be located significantly far (>40 m) away from adults, a result confirmed by direct testing in a second 56-ha plot. Sapling growth over 6 years, the distribution of newly emerging seedlings around adults, recruitment of saplings in a large opening and the outward extent of seedlings at the grove edge were also investigated. Two processes appear to have been operating: (1) a very strong and consistent restriction of the very numerous seedlings establishing after masting close to adults, and (2) a strong but highly spatially variable promotion of distant survivors by increased light from the deaths of large trees of species other than M. bisulcata (which itself has very low mortality rate). This leads to an apparent escape-from-adults effect. To maintain saplings in the shade between multiple short periods of release ectomycorrhizal connections to other co-occurring caesalp species may enable a rachet-type mechanism. The recorded sapling dynamics currently contribute an essential part of the long-term cycling of the groves. M. bisulcata is an interesting example of an important group of tropical trees, particularly in Africa, which are both highly light-demanding when young yet capable also of forming very large forest emergents. To more comprehensively explain tropical tree responses, the case is made for adding a new dimension to the trade-off concept of early tree light-response versus adult longevity.

Towards greater accuracy in individual-tree mortality regression

Background mortality is an essential component of any forest growth and yield model. Forecasts of mortality contribute largely to the variability and accuracy of model predictions at the tree, stand and forest level. In the present study, I implement and evaluate state-of-the-art techniques to increase the accuracy of individual tree mortality models, similar to those used in many of the current variants of the Forest Vegetation Simulator, using data from North Idaho and Montana. The first technique addresses methods to correct for bias induced by measurement error typically present in competition variables. The second implements survival regression and evaluates its performance against the traditional logistic regression approach. I selected the regression calibration (RC) algorithm as a good candidate for addressing the measurement error problem. Two logistic regression models for each species were fitted, one ignoring the measurement error, which is the “naïve” approach, and the other applying RC. The models fitted with RC outperformed the naïve models in terms of discrimination when the competition variable was found to be statistically significant. The effect of RC was more obvious where measurement error variance was large and for more shade-intolerant species. The process of model fitting and variable selection revealed that past emphasis on DBH as a predictor variable for mortality, while producing models with strong metrics of fit, may make models less generalizable. The evaluation of the error variance estimator developed by Stage and Wykoff (1998), and core to the implementation of RC, in different spatial patterns and diameter distributions, revealed that the Stage and Wykoff estimate notably overestimated the true variance in all simulated stands, but those that are clustered. Results show a systematic bias even when all the assumptions made by the authors are guaranteed. I argue that this is the result of the Poisson-based estimate ignoring the overlapping area of potential plots around a tree. Effects, especially in the application phase, of the variance estimate justify suggested future efforts of improving the accuracy of the variance estimate. The second technique implemented and evaluated is a survival regression model that accounts for the time dependent nature of variables, such as diameter and competition variables, and the interval-censored nature of data collected from remeasured plots. The performance of the model is compared with the traditional logistic regression model as a tool to predict individual tree mortality. Validation of both approaches shows that the survival regression approach discriminates better between dead and alive trees for all species. In conclusion, I showed that the proposed techniques do increase the accuracy of individual tree mortality models, and are a promising first step towards the next generation of background mortality models. I have also identified the next steps to undertake in order to advance mortality models further.

Herbivores equalize the seedling height growth of three dominant tree species in an African tropical rain forest

Determining the impact of insect herbivores on forest tree seedlings and saplings is difficult without experimentation in the field. Moreover, this impact may be heterogeneous in time and space because of seasonal rainfall and canopy disturbances, or ‘gaps’, which can influence both insect abundance and plant performance. In this study we used fine netting to individually protect seedlings of Microberlinia bisulcata, Tetraberlinia bifoliolata and Tetraberlinia korupensis trees (Fabaceae = Leguminosae) from insects in 41 paired gap-understorey locations across 80 ha of primary rain forest (Korup, Cameroon). For all species, growth in height and leaf numbers was negligible in the understorey, where M. bisulcata had the lowest survival after c. 2 years. In gaps, however, all species responded positively with pronounced above-ground growth across seasons. When exposed to herbivores their seedling height growth was similar, but in the absence of herbivores, M. bisulcata significantly outgrew both Tetraberlinia species and matched their leaf numbers. This result suggests that insect herbivores might play an important role in maintaining species coexistence by mitigating sapling abundance of the more palatable M. bisulcata, which in gaps was eaten the most severely. The higher ratio in static leaf damage of control-to-caged M. bisulcata seedlings in gaps than understorey locations was consistent with the Plant Vigour Hypothesis. This result, however, did not apply to either Tetraberlinia species. For M. bisulcata and T. korupensis, but not T. bifoliolata (the most shade-tolerant species), caging improved relative seedling survival in the understory locations compared to gaps, providing restricted support for the Limiting Resource Model. Approximately 2.25 years after treatments were removed, the caged seedlings were taller and had more leaves than controls in all three species, and the effect remained strongest for M. bisulcata. We conclude that in this community the impact of leaf herbivory on seedling growth in gaps is strong for the dominant M. bisulcata, which coupled to a very low shade-tolerance contributes to limiting its regeneration. However, because gaps are common to most forests, insect herbivores may be having impacts upon functionally similar tree species that are also characterized by low sapling recruitment much more widely than currently appreciated. An implication for the restoration and management of M. bisulcata populations in forests outside of Korup is that physical protection from herbivores of new seedlings where the canopy is opened by gaps, or by harvesting, should substantially increase its subcanopy regeneration, and thus, too, its opportunities for adult recruitment.

Carbon stocks, tree diversity, and the role of organic certification in different cocoa production systems in Alto Beni, Bolivia

This study compares aboveground and belowground carbon stocks and tree diversity in different cocoa cultivation systems in Bolivia: monoculture, simple agroforestry, and successional agroforestry, as well as fallow as a control. Since diversified, agroforestry-based cultivation systems are often considered important for sustainable development, we also evaluated the links between carbon stocks and tree diversity, as well as the role of organic certification in transitioning from monoculture to agroforestry. Biomass, tree diversity, and soil physiochemical parameters were sampled in 15 plots measuring 48 × 48 m. Semi-structured interviews with 52 cocoa farmers were used to evaluate the role of organic certification and farmers’ organizations (e.g., cocoa cooperatives) in promoting tree diversity. Total carbon stocks in simple agroforestry systems (128.4 ± 20 Mg ha−1) were similar to those on fallow plots (125.2 ± 10 Mg ha−1). Successional agroforestry systems had the highest carbon stocks (143.7 ± 5.3 Mg ha−1). Monocultures stored significantly less carbon than all other systems (86.3 ± 4.0 Mg ha−1, posterior probability P(Diff > 0) of 0.000–0.006). Among shade tree species, Schizolobium amazonicum, Centrolobium ochroxylum, and Anadenanthera sp. accumulated the most biomass. High-value timber species (S. amazonicum, C. ochroxylum, Amburana cearensis, and Swietenia macrophylla) accounted for 22.0 % of shade tree biomass. The Shannon index and tree species richness were highest in successional agroforestry systems. Cocoa plots on certified organic farms displayed significantly higher tree species richness than plots on non-certified farms. Thus, expanding the coverage of organic farmers’ organizations may be an effective strategy for fostering transitions from monoculture to agroforestry systems.

Seedling resistance, tolerance and escape from herbivores: insights from co-dominant canopy tree species in a resource-poor African rain forest

* Although plants can reduce the impacts of herbivory in multiple ways, these defensive traits are often studied in isolation and an understanding of the resulting strategies is incomplete. * In the study reported here, empirical evidence was simultaneously evaluated for the three main sets of traits available to plants: (i) resistance through constitutive leaf traits, (ii) tolerance to defoliation and (iii) escape in space, for three caesalpiniaceous tree species Microberlinia bisulcata, Tetraberlinia bifoliolata and T. korupensis, which co-dominate groves within the lowland primary rain forest of Korup National Park (Cameroon). * Mesh cages were placed around individual wild seedlings to exclude insect herbivores at 41 paired canopy gap and understorey locations. After following seedling growth and survival for c. 2 years, caged and control treatments were removed, leaves harvested to determine nutrient and phenolic concentrations, leaf mass per area estimated, and seedling performance in gaps followed for a further c. 2 years to quantify tolerance to the leaf harvesting. * The more nutrient-rich leaves of the weakly shade-tolerant M. bisulcata were damaged much more in gaps than the two strongly shade-tolerant Tetraberlinia species, which had higher leaf mass per area and concentrations of total phenols. Conversely, the faster-growing M. bisulcata was better able to tolerate defoliation in terms of height growth (reflushing capacity), but not at maintaining overall leaf numbers, than the other two species. * Across gaps, insect-mediated Janzen–Connell effects were most pronounced for M. bisulcata, less so for T. korupensis, and not detectable for T. bifoliolata. The three species differed distinctly in their secondary metabolic profiles. * Taken together, the results suggested a conceptual framework linking the three sets of traits, one in which the three co-dominant species adopt different strategies towards herbivore pressure depending on their different responses to light availability. This study is one of the first in a natural forest ecosystem to examine resistance to, tolerance of, and escape from herbivory among a group of co-occurring tropical tree species.

Limitation of seedling growth by potassium and magnesium supply for two ectomycorrhizal tree species of a Central African rain forest and its implication for their recruitment

In the ectomycorrhizal caesalpiniaceous groves of southern Korup National Park, the dominant tree species, Microberlinia bisulcata, displays very poor in situ recruitment compared with its codominant, Tetraberlinia bifoliolata. The reported ex situ experiment tested whether availabilities of soil potassium and magnesium play a role. Seedlings of the two species received applications of K and Mg fertilizer in potted native soil in a local shade house, and their responses in terms of growth and nutrient concentrations were recorded over 2 years. Amended soil concentrations were also determined. Microberlinia responded strongly and positively in its growth to Mg, but less to K; Tetraberlinia responded weakly to both. Added Mg led to strongly increased Mg concentration for Microberlinia while added K changed that concentration only slightly; Tetraberlinia strongly increased its concentration of K with added K, but only somewhat its Mg concentration with added Mg. Additions of Mg and K had small but important antagonistic effects. Microberlinia is Mg-demanding and apparently Mg-limited in Korup soil; Tetraberlinia, whilst K-demanding, appeared not to be K-limited (for growth). Added K enhanced plant P concentrations of both species. Extra applied Mg may also be alleviating soil aluminum toxicity, and hence improving growth indirectly and especially to the benefit of Microberlinia. Mg appears to be essential for Microberlinia seedling growth and its low soil availability in grove soils at Korup may be an important contributing factor to its poor recruitment. Microberlinia is highly shade-intolerant and strongly light-responding, whilst Tetraberlinia is more shade-tolerant and moderately light-responding, which affords an interesting contrast with respect to their differing responses to Mg supply. The study revealed novel aspects of functional traits and likely niche-partitioning among ectomycorrhizal caesalps in African rain forests. Identifying the direct and interacting indirect effects of essential elements on tropical tree seedling growth presents a considerable challenge due the complex nexus of causes involved.

Canopy gaps promote selective stem-cutting by small mammals of two dominant tree species in an African lowland forest: the importance of seedling chemistry

Small mammals can impede tree regeneration by injuring seedlings and saplings in several ways. One fatal way is by severing their stems, but apparently this type of predation is not well-studied in tropical rain forest. Here, we report on the incidence of 'stem-cutting' to new, wild seedlings of two locally dominant, canopy tree species monitored in 40 paired forest understorey and gap-habitat areas in Korup, Cameroon following a 2007 masting event. In gap areas, which are required for the upward growth and sapling recruitment of both species, 137 seedlings of the long-lived, light-demanding, fast-growing large tropical tree (Microberlinia bisulcata) were highly susceptible to stem-cutting (83% of deaths) - it killed 39% of all seedlings over a c. 2-y period. In stark contrast, seedlings of the more shade-tolerant, slower-growing tree species (Tetraberlinia bifoliolata) were hardly attacked (4.3%). In the understorey, however, stem-cutting was virtually absent. Across the gap areas, the incidence of stem-cutting of M. bisulcata seedlings showed significant spatial variation that could not be explained significantly by either canopy openness or Janzen-Connell type effects (proximity and basal area of conspecific adult trees). To examine physical and chemical traits that might explain the species difference to being cut, bark and wood tissues were collected from a separate sample of seedlings in gaps (i.e. not monitored for stem-cutting). These analyses suggested that, compared with T. bifoliolata, the lower stem density, higher Mg and K and fatty acid concentrations in bark, and fewer phenolic and terpene compounds in M. bisulcata seedlings made them more palatable and attractive to small-mammal predators, likely rodents. We conclude that selective stem-cutting is a potent countervailing force to the current local canopy dominance of the grove-forming M. bisulcata by limiting the recruitment and abundance of its saplings. Given the ubiquity of gaps and ground-dwelling rodents in pantropical forests, it would be surprising if this form of lethal browsing was restricted to Korup.

Light and seed size affect establishment of grove-forming ectomycorrhizal rain forest tree species

• Regeneration of the dominant ectomycorrhizal tree Microberlinia bisulcata in groves in Korup, Central Africa, is very poor. The hypothesis was tested that this species is more shade intolerant than other co-occurring species. • In two 1-yr trials, each with M. bisulcata and four other species at a nursery close to Korup, growth was measured under five PAR levels, with ± added P and ± watering in the dry season. In parallel experiments the effects of PAR with two R : FR ratios were investigated. • Increasing PAR had a consistent effect on the rates of increase in plant mass and on changes in the other variables. Doubling soil P, watering and halving the R : FR ratio had almost no effect. However, across species, mass at low PAR and relative growth rate related positively and negatively, respectively, to seed mass. • One contributing factor for the poor recruitment of M. bisulcata is therefore its low survival and slow growth at low PAR, due to its small seed size. The two codominant ectomycorrhizal grove species of Tetraberlinia, with larger seeds, were less affected by low PAR.