L’arbre de régression multivariable et les modèles linéaires généralisés revisités : applications à l’étude de la diversité bêta et à l’estimation de la biomasse d’arbres tropicaux

En écologie, dans le cadre par exemple d’études des services fournis par les écosystèmes, les modélisations descriptive, explicative et prédictive ont toutes trois leur place distincte. Certaines situations bien précises requièrent soit l’un soit l’autre de ces types de modélisation ; le bon choix s’impose afin de pouvoir faire du modèle un usage conforme aux objectifs de l’étude. Dans le cadre de ce travail, nous explorons dans un premier temps le pouvoir explicatif de l’arbre de régression multivariable (ARM). Cette méthode de modélisation est basée sur un algorithme récursif de bipartition et une méthode de rééchantillonage permettant l’élagage du modèle final, qui est un arbre, afin d’obtenir le modèle produisant les meilleures prédictions. Cette analyse asymétrique à deux tableaux permet l’obtention de groupes homogènes d’objets du tableau réponse, les divisions entre les groupes correspondant à des points de coupure des variables du tableau explicatif marquant les changements les plus abrupts de la réponse. Nous démontrons qu’afin de calculer le pouvoir explicatif de l’ARM, on doit définir un coefficient de détermination ajusté dans lequel les degrés de liberté du modèle sont estimés à l’aide d’un algorithme. Cette estimation du coefficient de détermination de la population est pratiquement non biaisée. Puisque l’ARM sous-tend des prémisses de discontinuité alors que l’analyse canonique de redondance (ACR) modélise des gradients linéaires continus, la comparaison de leur pouvoir explicatif respectif permet entre autres de distinguer quel type de patron la réponse suit en fonction des variables explicatives. La comparaison du pouvoir explicatif entre l’ACR et l’ARM a été motivée par l’utilisation extensive de l’ACR afin d’étudier la diversité bêta. Toujours dans une optique explicative, nous définissons une nouvelle procédure appelée l’arbre de régression multivariable en cascade (ARMC) qui permet de construire un modèle tout en imposant un ordre hiérarchique aux hypothèses à l’étude. Cette nouvelle procédure permet d’entreprendre l’étude de l’effet hiérarchisé de deux jeux de variables explicatives, principal et subordonné, puis de calculer leur pouvoir explicatif. L’interprétation du modèle final se fait comme dans une MANOVA hiérarchique. On peut trouver dans les résultats de cette analyse des informations supplémentaires quant aux liens qui existent entre la réponse et les variables explicatives, par exemple des interactions entres les deux jeux explicatifs qui n’étaient pas mises en évidence par l’analyse ARM usuelle. D’autre part, on étudie le pouvoir prédictif des modèles linéaires généralisés en modélisant la biomasse de différentes espèces d’arbre tropicaux en fonction de certaines de leurs mesures allométriques. Plus particulièrement, nous examinons la capacité des structures d’erreur gaussienne et gamma à fournir les prédictions les plus précises. Nous montrons que pour une espèce en particulier, le pouvoir prédictif d’un modèle faisant usage de la structure d’erreur gamma est supérieur. Cette étude s’insère dans un cadre pratique et se veut un exemple pour les gestionnaires voulant estimer précisément la capture du carbone par des plantations d’arbres tropicaux. Nos conclusions pourraient faire partie intégrante d’un programme de réduction des émissions de carbone par les changements d’utilisation des terres.

Biomass and mineralmass estimates in a "cerrado" ecosystem

This work aimed to develop allometric equations for tree biomass estimation, and to determine the site biomass in different "cerrado" ecosystems. Destructive sampling in a "campo cerrado" (open savanna) was carried out at the Biological Reserve of Moji-Guaçu, State of São Paulo, southeastern Brazil. This "campo cerrado" (open savanna) grows under a tropical climate and on acid, low nutrient soils. Sixty wood plants were cut to ground level and measurements of diameter, height and weight of leaves and stems were taken. We selected the best equations among the most commonly used mathematical relations according to R² values, significance, and standard error. Both diameter (D) and height (H) showed good relationship with plant biomass, but the use of these two parameters together (DH and D²H) provided the best predictor variables. The best equations were linear, but power and exponential equations also showed high R² and significance. The applicability of these equations is discussed and biomass estimates are compared with other types of tropical savannas. Mineralmass was also estimated. "Cerrados" proved to have very important carbon reservoirs due to their great extent. In addition, high land-use change that takes place nowadays in the "cerrado" biome may significantly affect the global carbon cycle.

Leaf synchrony and insect herbivory among tropical tree habitat specialists

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Tree height integrated into pantropical forest biomass estimates

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- and Weibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (< 40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8 Mg ha(-1) (range 6.6 to 112.4) to 8.0 Mg ha(-1) (-2.5 to 23.0).

Tree height integrated into pantropical forest biomass estimates

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- andWeibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (?40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8Mgha?1 (range 6.6 to 112.4) to 8.0Mgha?1 (?2.5 to 23.0). For all plots, aboveground live biomass was ?52.2 Mgha?1 (?82.0 to ?20.3 bootstrapped 95%CI), or 13%, lower when including H estimates, with the greatest relative reductions in estimated biomass in forests of the Brazilian Shield, east Africa, and Australia, and relatively little change in the Guiana Shield, central Africa and southeast Asia. Appreciably different stand structure was observed among regions across the tropical continents, with some storing significantly more biomass in small diameter stems, which affects selection of the best height models to reduce uncertainty and biomass reductions due to H. After accounting for variation in H, total biomass per hectare is greatest in Australia, the Guiana Shield, Asia, central and east Africa, and lowest in eastcentral Amazonia, W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if tropical forests span 1668 million km2 and store 285 Pg C (estimate including H), then applying our regional relationships implies that carbon storage is overestimated by 35 PgC (31?39 bootstrapped 95%CI) if H is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree H is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of tropical carbon stocks and emissions due to deforestation.

Umbrella pine above ground biomass estimation with very high spatial resolution satellite images

Above ground biomass is frequently estimated with forest inventory data and an extrapolation method for the per unit area evaluations. This procedure is labour demanding and costly. In this study above ground biomass functions, whose independent variable is crown horizontal projection, were developed. Multi-resolution segmentation method and object-oriented classification, based on very high spatial resolution satellite images, were used to obtain the area of tree crown horizontal projection for umbrella pine (Pinus pinea L.). A set of inventory plots were measured and with existing allometric functions for this species above ground biomass per tree and per plot were calculated. The two data sets were used to fit linear functions both for individual plot and their cumulative values. The results show a good performance of the models. Errors smaller than 10% are obtained for stand areas greater than 1.4 ha. These functions have the advantages of estimating above ground biomass for all the area under study or surveillance, not requiring forest inventory; allow monitoring in short time periods; and are easily implemented in a geographical information system environment.

Above ground biomass estimation in Mediterranean multiple use systems with high spatial resolution satellite images

Forest biomass has been having an increasing importance in the world economy and in the evaluation of the forests development and monitoring. It was identified as a global strategic reserve, due to its applications in bioenergy, bioproduct development and issues related to reducing greenhouse gas emissions. The estimation of above ground biomass is frequently done with allometric functions per species with plot inventory data. An adequate sampling design and intensity for an error threshold is required. The estimation per unit area is done using an extrapolation method. This procedure is labour demanding and costly. The mail goal of this study is the development of allometric functions for the estimation of above ground biomass with ground cover as independent variable, for forest areas of holm aok (Quercus rotundifolia), cork oak (Quercus suber) and umbrella pine (Pinus pinea) in multiple use systems. Ground cover per species was derived from crown horizontal projection obtained by processing high resolution satellite images, orthorectified, geometrically and atmospheric corrected, with multi-resolution segmentation method and object oriented classification. Forest inventory data were used to estimate plot above ground biomass with published allometric functions at tree level. The developed functions were fitted for monospecies stands and for multispecies stands of Quercus rotundifolia and Quercus suber, and Quercus suber and Pinus pinea. The stand composition was considered adding dummy variables to distinguish monospecies from multispecies stands. The models showed a good performance. Noteworthy is that the dummy variables, reflecting the differences between species, originated improvements in the models. Significant differences were found for above ground biomass estimation with the functions with and without the dummy variables. An error threshold of 10% corresponds to stand areas of about 40 ha. This method enables the overall area evaluation, not requiring extrapolation procedures, for the three species, which occur frequently in multispecies stands.

Reserve carbohydrates and lipids from the seeds of four tropical tree species with different sensitivity to desiccation

Considering the importance of water content for the conservation and storage of seeds, and the involvement of soluble carbohydrates and lipids for embryo development, a comparative study was carried out among the seeds of Inga vera (ingá), Eugenia uniflora (pitanga), both classified as recalcitrant, and Caesalpinia echinata (brazilwood) and Erythrina speciosa (mulungu), considered as orthodox seeds. Low concentrations of cyclitols (0.3-0.5%), raffinose family oligosaccharides (ca. 0.05%) and unsaturated fatty acids (0-19%) were found in the seeds of ingá and pitanga, while larger amounts of cyclitols (2-3%) and raffinose (4.6-13%) were found in brazilwood and mulungu, respectively. These results, in addition to higher proportions of unsaturated fatty acids (53-71%) in orthodox seeds, suggested that sugars and lipids played important role in water movement, protecting the embryo cell membranes against injuries during dehydration.

Tropical woods: influence of density in hardness parallel and normal to the grain for some Brazilian tropical tree species

The aim of this work is to verify the possibility to correlating specific gravity and wood hardness parallel and perpendicular to the grain. The purpose is to offer one more tool to help in the decision about wood species choice for use in floors and sleepers. To reach this intent, we considered the results of standard tests (NBR 7190:1997, Timber Structures Design, Annex B, Brazilian Association of Technical Standards) to determine hardness parallel and normal to the grain in fourteen tropical high density wood species (over 850 kg/m(3), at 12% moisture content). For each species twelve determinations were made, based on the material obtained at Sao Carlos and its regional wood market. Statistical analysis led to some expressions to describe the cited properties relationships, with a determination coefficient about 0.8.

Differential responses of antioxidant enzymes in pioneer and late-successional tropical tree species grown under sun and shade conditions

To investigate the ability of pioneer and late-successional species to adapt to a strong light environment in a reforestation area, we examined the activities of antioxidant enzymes in relation to photosystem chlorophyll a fluorescence and photosynthetic pigment concentration for eight tropical tree species grown under 100% (sun) and 10% (shade) sunlight irradiation. The pioneer (early-succession) species (PS) were Cecropia pachystachya, Croton urucurana, Croton floribundus and Schinus terebinthifolius. The non-pioneer (late succession) species (LS) were Hymenaea courbaril L var. stilbocarpa, Esenbeckia leiocarpa, Cariniana legalis and Tabebuia roseo-alba. We observed a greater decline in the ratio of variable to maximum chlorophyll a fluorescence (F(v)/F(m)) under full sunlight irradiation in the late-successional species than in the pioneer species. The LS species most sensitive to high irradiance were C. legalis and H. courbaril. In LS species, chlorophyll a, chlorophyll b and total chlorophyll concentrations were higher in the shade-grown plants than in plants that developed under full sunlight, but in the PS species C. floribundus and C. pachystachya, we did not observe significant changes in chlorophyll content when grown in the two contrasting environments. The carotenoids/total chlorophyll ratio increased significantly when plants developed under high-sunlight irradiation, but this response was not observed in the PS species S. terebinthifolius and C. pachystachya. The improved performance of the pioneer species in high sunlight was accompanied by an increase in superoxide dismutase (SOD. EC 1.15.1.1) activity, though no light-dependent increase in the activity of ascorbate peroxidase (APX. EC 1.11.1.11) was observed. The activity of catalase (CAT, EC 1.11.1.6) was reduced by high irradiation in both pioneer and late-successional species. Our results show that pioneer species perform better under high-sunlight irradiation than late-successional species, as indicated by increased SOD activity and a higher F IF,, ratio. C. legalis was the LS species most susceptible to photoinhibition under full sunlight conditions. These results suggest that pioneer plants have more potential tolerance to photo-oxidative damage than late-successional species associated with the higher SOD activity found in pioneer species. Reduced photoinhibition in pioneer species probably results from their higher photosynthetic capacities, as has been observed in a previous survey carried out by our group. (C) 2010 Elsevier B.V. All rights reserved.

Microsatellite diversity and effective population size in a germplasm bank of Hymenaea courbaril var. stilbocarpa (Leguminosae), an endangered tropical tree: recommendations for conservation

Deforestation in southeast Brazil has led to the extinction of Hymenaea courbaril var. stilbocarpa and ex situ conservation has been established. In this study, the levels of genetic diversity and the effective population size of H. courbaril in a germplasm bank were investigated using six nuclear microsatellite loci. A total of 79 and 91 alleles were found in 65 seed-trees and their 176 offspring, respectively. Offspring have a higher average number of alleles per locus (A = 15.2) than seed-trees (A = 13.2), but lower observed heterozygosity (offspring: H (o) = 0.566; seed-trees: H (o) = 0.607). The estimate of outcrossing rate shows that the study population is perfectly outcrossed (t (m) = 0.978, P > 0.05). Significant deviations from random mating were detected through mating among relatives and correlated matings. The average variance in effective population size for each family was 2.63, with a total effective population size retained in the bank of 170.1. These results confirm that the preserved population of H. courbaril retains substantial genetic variability.

Development and characterization of 15 microsatellite loci for Cariniana estrellensis and transferability to Cariniana legalis, two endangered tropical tree species

From a genomic library enriched for GA/CA repeats, 15 highly polymorphic microsatellite markers were developed for Cariniana estrellensis, a tropical forest tree. The microsatellite loci were screened in 49 mature trees found between Pardo river and Mogi-Gua double dagger u river basins, in the state of So Paulo, Brazil. A total of 140 alleles were detected with an average of 9.33 alleles per locus. The expected heterozygosity ranged from 0.37 to 0.88. These loci showed a high probability of paternity exclusion. Additionally, 12 loci were effectively transferred to Cariniana legalis. High levels of polymorphism make the present SSR markers useful for population genetic studies.

Transferability and characterization of nine microsatellite markers for the tropical tree species Tabebuia roseo-alba

Microsatellite loci that were previously developed in the tropical tree Tabebuia aurea were used for the genetic analysis of Tabebuia roseo-alba populations. Nine of 10 simple sequence repeat markers were amplified, and the polymorphism was assessed in 58 individuals sampled from two stands in southeastern Brazil. All loci were polymorphic with Mendelian inheritance. The allele numbers were high, ranging from 5 to 13 in population I and 3 to 7 in population II, with means of 8.9 and 5.5, respectively. We conclude that these markers can be efficiently used for parentage and gene-flow studies.

Exploring TM image texture and its relationships with biomass estimation in Rondônia, Brazilian Amazon

Many texture measures have been developed and used for improving land-cover classification accuracy, but rarely has research examined the role of textures in improving the performance of aboveground biomass estimations. The relationship between texture and biomass is poorly understood. This paper used Landsat Thematic Mapper (TM) data to explore relationships between TM image textures and aboveground biomass in Rondônia, Brazilian Amazon. Eight grey level co-occurrence matrix (GLCM) based texture measures (i.e., mean, variance, homogeneity, contrast, dissimilarity, entropy, second moment, and correlation), associated with seven different window sizes (5x5, 7x7, 9x9, 11x11, 15x15, 19x19, and 25x25), and five TM bands (TM 2, 3, 4, 5, and 7) were analyzed. Pearson's correlation coefficient was used to analyze texture and biomass relationships. This research indicates that most textures are weakly correlated with successional vegetation biomass, but some textures are significantly correlated with mature forest biomass. In contrast, TM spectral signatures are significantly correlated with successional vegetation biomass, but weakly correlated with mature forest biomass. Our findings imply that textures may be critical in improving mature forest biomass estimation, but relatively less important for successional vegetation biomass estimation.