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The loss of biodiversity is affecting all ecosystems on Earth, one of the greatest threats to biodiversity being climate change. Forests have been highlighted for the potential to mitigate climate change by storing carbon above- and belowground in soils. In this thesis, I studied the effects of tree diversity on carbon cycling in subtropical Chinese forests. I aimed to explore the mechanisms behind tree diversity effects on carbon cycling by focusing on microbial-based processes and the consequences of tree diversity-induced spatial heterogeneity. First, my colleagues and I tested the effects of tree diversity on litterfall spatial patterns and the consequences for litter decomposition and quantified the importance of microbial community in decomposition processes. Second, we explored the effects of tree diversity on relationships between soil microbial facets and soil microbial functions. Third, we tested the effects of tree diversity on soil microbial biomass and carbon concentrations, and their mediation by biotic and abiotic conditions. Finally, we explored the consequences of diversifying forests for re-/afforestation initiatives and plantations to reduce atmospheric carbon levels, and the benefits of diversity for mitigating the effects of climate change on ecosystems and human well-being. We highlighted the positive effects of tree diversity on tree productivity. By increasing the amount and diversity of litterfall, tree diversity increased litter decomposition and subsequently the assimilation of tree products into the forest soils. Our investigation has shown the key role of microbial communities for forests carbon dynamics by carrying out litter decomposition, soil heterotrophic respiration, and soil carbon stabilization. Most notably, tree diversity effects on soil microbial respiration were mainly mediated by soil microbial biomass rather than soil microbial community taxonomic or functional diversity. The effects of tree diversity on microbial biomass were mediated by biotic and abiotic conditions. Taken together, we revealed the importance of considering space to understand biodiversity-ecosystem functioning relationships. Finally, we argued that tree diversity is a promising avenue to maximize the potential of re-/afforestation projects to mitigate increasing atmospheric carbon. Moreover, we highlighted that diversifying forests in re-/afforestation initiatives can help to reduce climate change effects on ecosystems: first, by increasing resistance and resilience to extreme climatic events, and second, by buffering microclimatic conditions in natural and urban areas. My investigation highlighted that tree diversity effects on ecosystem functioning could be explained by both mass and diversity effects on higher trophic levels and their functions. In addition, I showed the key role of tree diversity-induced spatial heterogeneity and the need to consider space and time in further research. Moreover, these results need to be combined with practitioner constraints to enable feasible restoration projects.:Summary table Bibliographic information .................................................................................... I ~ XV Main body ......................................................................................................... 1 ~ 212 Supplementary materials ..................................................................................... i ~ xv Scientific supplementary materials ............................................................. -1- ~ - 154- Table of Contents Table of figures .......................................................................................................... XI Table of scientific supplementary materials ............................................................. XIII Glossary ................................................................................................................... XV Introduction ................................................................................................................. 3 Chapter I - Tree diversity effects on litter decomposition are mediated by litterfall and microbial processes .................................................................................................. 35 Transition I - II ........................................................................................................... 67 Chapter II - Tree diversity and soil chemical properties drive the linkages between soil microbial community and ecosystem functioning................................................ 71 Transition II - III ....................................................................................................... 107 Chapter III - Abiotic and biotic drivers of scale-dependent tree trait effects on soil microbial biomass and soil carbon concentration ................................................... 111 Transition III - IV ..................................................................................................... 155 Chapter IV – Diverse forests are cool: promoting diverse forests to mitigate carbon emissions and climate change ............................................................................... 159 General discussion ................................................................................................. 173 Abstract .................................................................................................................. 195 General acknowledgments ..................................................................................... 209 Supplementary materials ..............................................................................................i

The increasing frequency and intensity of droughts threaten forests and their climate change mitigation potential. Mixed-species forests are promoted to increase forest productivity and stability compared to monospecific forests, but we still lack a mechanistic understanding of the strength, nature and drivers of tree diversity effects on productivity and stability under drought. Here, I studied the stress hotter droughts inflict on trees and examined whether diversification in tree species, structures and drought-tolerance traits is a potential solution to this threat. In study 1, I found that the hotter drought years 2018–2019, the severest droughts since records, induced unprecedented tree productivity and physiological stress responses (reduced growth and increased δ13C) in a Central European floodplain forest. Hotter droughts thus constitute a novel threat. In studies 2–4, I examined diversity-productivity and diversity-stability relationships across spatiotemporal scales in a tropical (study 2) and a subtropical (studies 3, 4) tree diversity experiment specifically designed to examine biodiversity-ecosystem functioning relationships. Tree species richness consistently increased productivity and stability, and this effect was strongest at the highest levels of diversity. Structural diversity increased productivity but was unrelated to stability, while diversity in drought-tolerance traits increased stability but not productivity. Assessing drought-tolerance traits was essential for understanding the role of tree diversity during drought. Positive diversity effects on productivity scaled up from the tree neighbourhood to the community level, but effects on stability emerged only at the community level. Community stability increased with species richness due to asynchronous species responses to dry and wet years driven by species’ drought-tolerance traits. I showed that diversity but not identity in drought-tolerance traits increases community stability. Overall, promoting structurally and functionally diverse mixed-species forests may enable high productivity and stability under intensifying climate change.:1. General introduction 1.1. Mixed-species forests 1.2. Diversity-productivity relationships 1.3. Diversity-productivity relationships during drought 1.4. Diversity-stability relationships 1.5. Diversity facets 1.6. Drought-tolerance traits 1.7. Linkages between the four studies 2. Methodological features 2.1. Study sites and approaches 2.2. Productivity, stability and physiological water stress 2.3. The quantification of diversity 2.4. Spatiotemporal analyses 3. Original contributions Study 1: Cumulative growth and stress responses to the 2018–2019 drought in a European floodplain forest Study 2: Drivers of productivity and its temporal stability in a tropical tree diversity experiment Study 3: Neighbourhood species richness and drought-tolerance traits modulate tree growth and δ13C responses to drought Study 4: Species richness stabilizes productivity via asynchrony and drought- tolerance diversity in a large-scale tree biodiversity experiment 4. General discussion 4.1. Summary of main findings 4.2. Hotter droughts and forest functioning 4.3. Diversity signals across spatial scales 4.4. Diversity signals across temporal scales 4.5. Diversity facets 4.6. Context dependency and transferability 4.7. Implications for forest management in the 21st century 5. Outlook and future research 5.1. Observation and experimentation under hotter droughts 5.2. Response variables 5.3. Diversity facets 5.4. Drought-tolerance traits 5.5. Zooming in 5.6. Zooming out 5.7. From understanding to use of BEF relationships 6. Conclusion 7. Summary 8. Zusammenfassung 9. References Acknowledgements Author contribution statements Curriculum vitae List of publications Selbstständigkeitserklärung

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Résumé: Deux facteurs principaux, une fois combinés, permettent de comprendre l’assemblage des communautés, soient i) l’environnement (abiotique et biotique), qui agit comme un filtre sélectionnant les espèces les mieux adaptées aux conditions données, et ii) les traits fonctionnels, sur lesquels s’effectue ce filtrage environnemental puisqu’ils représentent les adaptations des espèces aux conditions données. Il est donc essentiel d’établir des relations fiables entre les conditions environnementales et la structure fonctionnelle des communautés afin de pouvoir identifier et comprendre les mécanismes régissant l’assemblage des communautés. Cependant, plusieurs facteurs tels que les interactions entre variables environnementales à différentes échelles spatiales (par exemple le climat et la fertilité des sols) peuvent complexifier la situation et c’est pourquoi, malgré une quantité grandissante d’études sur le sujet, les processus d’assemblage des communautés restent difficiles à définir et à généraliser. Cette thèse vise donc à i) mieux définir et quantifier les relations trait-environnement des systèmes d’herbacées au travers de différentes échelles spatiales et ii) déterminer l’influence de ces relations sur l’assemblage des communautés et le fonctionnement des écosystèmes. Pour réaliser ces objectifs, j’ai travaillé avec les données de programmes ayant collecté des données taxonomiques et fonctionnelles sur les communautés d’herbacées à travers la France (DivHerbe et DivGrass) et, à moindre mesure, l’Europe (VISTA). Ces bases de données couvrent donc de larges gradients climatiques régionaux ainsi que des gradients environnementaux plus locaux relatifs à la qualité des sols et aux perturbations. J’ai d’abord testé l’importance de considérer à la fois des variables environnementales locales et régionales ainsi que leurs interactions pour déterminer la structure fonctionnelle et taxonomique des communautés. J’ai ensuite étudié comment l’importance relative des processus menant soit à la convergence ou à la divergence fonctionnelle peut changer le long d’une succession, puis comment ces deux types de processus influencent notre capacité à prédire l’assemblage des communautés à partir des traits fonctionnels. Finalement, j’ai présenté comment les résultats au niveau des communautés peuvent être utiles pour étudier le niveau des écosystèmes. Les résultats de cette thèse démontrent que les variables climatiques régionales interagissent fortement avec les variables environnementales locales pour influencer les processus locaux déterminant l’assemblage des communautés. Évaluer le contexte régional semble donc nécessaire afin d’éviter des interprétations erronées des patrons d’assemblage observés. Travaillant avec ces deux niveaux de variation environnementale, une dissociation important entre la variation taxonomique et fonctionnelle des communautés a été mise à jour, reflétant l’importance de considérer plusieurs facettes de biodiversité pour comprendre la dynamique des communautés. Les résultats ont aussi démontré que les processus d’assemblage menant à la convergence et à la divergence ont un impact très différent et prédictible sur les relations liant les traits et les abondances des espèces, c’est-à-dire notre capacité à prédire l’assemblage des communautés à partir des traits. Finalement, toutes ces notions, relatives aux relations trait-environnement et à l’assemblage des communautés basé sur les traits, ont été utilisées dans un contexte de biogéographie fonctionnelle. Il a été démontré qu’il était possible de construire des cartes de valeurs de traits fonctionnels dans les prairies permanentes à l’échelle de la France, à partir de variables environnementales. Certaines propriétés écosystémiques ont ensuite pu être prédites à partir de ces cartes. Ces travaux ont donc permis d’illustrer les défis à surmonter pour utiliser nos connaissances de l’écologie fonctionnelle en vue d’une conservation et d’une exploitation viables de nos écosystèmes. / Abstract: There are two main factors which, combined together, allow understanding community assembly : i) the environment (both abiotic and biotic), which acts as a filter selecting species according to how well-adapted they are to given conditions, and ii) functional traits, on which this environmental filtering occurs since they represent species adaptations to particular conditions. It is thus essential to establish reliable relationships between environmental conditions and the functional structure of communities in order to identify and understand the mechanisms driving community assembly. However, several factors such as cross-scale interactions between environmental variables (e.g. between climate and soil fertility) complicate the situation. This is why, despite a growing body of studies on the subject, processes of community assembly are still poorly understood and are difficult to generalize. The purpose of this thesis is to i) better define and quantify the trait-environment relationships in herbaceous systems across different spatial scales and ii) determine the influence of those relationships on community assembly and on ecosystem functioning. To realize these objectives, I worked with data from programs which assembled taxonomic and functional data on herbaceous communities across France (DivHerbe and DivGrass) and, to a lesser extent, Europe (VISTA). These databases thus cover large regional climatic gradients, as well as more local environmental gradients related to soil quality and disturbances. I first tested the importance of simultaneously considering local and regional environmental variables as well as their interactions to determine the taxonomic and functional structure of communities. Then, I studied how the relative importance of processes leading to either functional convergence or divergence can change along a successional gradient, and how these two types of processes influence our ability to predict community assembly from functional traits. Finally, I presented how the results at the community-level can be used to study the ecosystem-level. The results of this thesis demonstrate that regional climatic variables strongly interact with local environmental variables in driving the local processes responsible for community assembly. Assessing the regional context is thus necessary in order to avoid erroneous interpretations of observed assembly patterns. Working with those two levels of environmental variation, important discrepancies were found between taxonomic and functional variations across communities, reflecting the importance of considering several aspects of biodiversity in order to understand community dynamics. The results also demonstrated that the assembly processes leading to functional convergence and divergence have a very different and predictable impact on the relationships between traits and species relative abundances, i.e. on our ability to predict community assembly from traits. Finally, these notions related to trait-environment relationships and to trait-based community assembly were used in a functional biogeography framework: It was possible to build maps of functional traits values in permanent grasslands across France using environmental variables. These maps then allowed predicting particular ecosystem properties. Thus, this work allowed illustrating some challenges that we are facing in using our knowledge in functional ecology to build sustainable conservation and exploitation plans for our ecosystems.

Future global change scenarios predict a dramatic loss of biodiversity for many regions in the world, potentially reducing the resistance and resilience of ecosystem functions. Once before, during Plio-Pleistocene glaciations, harsher climatic conditions in Europe as compared to North America led to a more depauperate tree flora. Here we hypothesize that this climate driven species loss has also reduced functional diversity in Europe as compared to North America. We used variation in 26 traits for 154 North American and 66 European tree species and grid-based co-occurrences derived from distribution maps to compare functional diversity patterns of the two continents. First, we identified similar regions with respect to contemporary climate in the temperate zone of North America and Europe. Second, we compared the functional diversity of both continents and for the climatically similar subregions using the functional dispersion-index (FDis) and the functional richness index (FRic). Third, we accounted in these comparisons for grid-scale differences in species richness, and, fourth, investigated the associated trait spaces using dimensionality reduction. For gymnosperms we find similar functional diversity on both continents, whereas for angiosperms functional diversity is significantly greater in Europe than in North America. These results are consistent across different scales, for climatically similar regions and considering species richness patterns. We decomposed these differences in trait space occupation into differences in functional diversity vs. differences in functional identity. We show that climate-driven species loss on a continental scale might be decoupled from or at least not linearly related to changes in functional diversity. This might be important when analyzing the effects of climate-driven biodiversity change on ecosystem functioning.

Die großflächige Umwandlung von tropischen Wäldern hat zu dramatischen Verlusten von Biodiversität und assoziierten Ökosystemdienstleistungen und –funktionen geführt. Indonesien ist ein besonders schwerwiegendes Beispiel für den Verlust von Waldflächen und Biodiversität. Landumnutzungen, in starkem Maße durch den Anbau von Ölpalmen vorangetrieben, stellen eine erhebliche Bedrohung für die außergewöhnlich hohe Biodiversität des Landes dar. Landwirtschaftssysteme, wie Agroforstsysteme, können hingegen genutzt werden, um die Biodiversität und Ökosystemfunktionen in von Ölpalmen dominierten Landschaften zu erhöhen. In Regionen, in denen Ölpalmplantagen bereits die Landschaft dominieren, kann diese Erhöhung nur durch systematische Renaturierung erfolgen. Die zugrunde liegenden ökologischen und soziökonomischen Prozesse und damit verbundene Beschränkungen und Kompromisse von Renaturierungsmaßnahmen in von Ölpalmen dominierten Landschaften sind jedoch weitgehend unbekannt. Um diese Wissenslücke zu schließen, habe ich mit Kollegen aus Deutschland und Indonesien ein Langzeitexperiment zur Erhöhung der Biodiversität aufgebaut. Wir haben experimentell Bäume in Form von „Inseln“ in eine konventionelle Ölpalmplantage gepflanzt und hierbei systematisch die Flächengröße, das Diversitätslevel und die Artzusammensetzung variiert. Wir haben hierfür sechs multifunktionale heimische Baumarten ausgewählt. Auf der Fläche der Bauminseln haben wir einen Teil der Ölpalmen gefällt, um die Lichtverfügbarkeit für die gepflanzten Bäume durch eine reduzierte Ölpalmdichte zu erhöhen. In dieser Doktorarbeit stelle ich den Aufbau des Experiments vor und gebe einen breiten Einblick in anfängliche Auswirkungen des Experiments, indem ich ökologische Aspekte in Betracht ziehe, sowie Veränderungen hinsichtlich des Ernteertrags. Da die Zeit kurz nach der Pflanzung ein Nadelöhr für die Langzeitetablierung der Bäume darstellt, ist sie sehr kritisch, um die erwünschten Renaturierungserfolge in der Zukunft zu erzielen. Des Weiteren kann die Anfangszeit auch aus der Sicht der Landwirte eine entscheidende Hürde darstellen, da der Nutzen der Bauminseln erst lange Zeit nach ihrer Pflanzung entsteht. Zuerst beschreibe ich Umweltvariablen und biotische Charakteristika der den experimentellen Flächen assoziierten Vegetation, Invertebraten und Vögel vor der Errichtung des Experiments, sowie anfängliche Auswirkungen des Experiments auf die Fauna. Ein Jahr nach der Errichtung des Experiments hatten die Baumpflanzungen einen insgesamt positiven Effekt auf die Artengemeinschaften von Vögeln und Invertebraten in der Plantage. Die Größe der Bauminseln wirkte sich lediglich auf die Diversität und Abundanz von Invertebraten positiv aus, die somit auf kleinskalige Veränderungen reagierten. Ausgehend von diesen Ergebnissen erwarte ich einen weiteren Anstieg der Biodiversität und Ökosystemfunktionen in der Zukunft. Danach berichte ich über den Anwuchserfolg der Bäume und ermittle die wichtigsten Determinanten für den Wuchs und das Überleben der Bäume. Insgesamt sind die Bäume gut angewachsen, jedoch gab es große Unterschiede zwischen den Baumarten hinsichtlich relativer Höhenwachstums-, Dickenwachstums- sowie Überlebensraten. Arten mit hohen Wachstumsraten wiesen vorwiegend auch hohe Überlebensraten auf. Auf Versuchsflächenebene hatten Standortbedingungen, Flächengröße und Diversitätslevel der gepflanzten Bäume lediglich einen geringen Einfluss auf das mittlere Baumwachstum und die Überlebensrate. Auf Individuenebene habe ich signifikante Nachbarschaftseffekte festgestellt. Hohe benachbarte Bäume und die Distanz zu verbleibenden Ölpalmen begünstigten die Entwicklung der Bäume. Noch ist es zu früh zu entscheiden, welche der Baumarten sich generell am besten eignen, um Renaturierungsziele zu erreichen. Unterschiede in der anfänglichen Entwicklung liefern jedoch wichtige Informationen für eine zukünftige Bewertung der Arten. Zuletzt präsentiere ich Auswirkungen des Experiments auf den Ölpalmertrag. Nach zwei Jahren waren die Erträge pro Ölpalmindividuum erhöht, sowohl auf als auch direkt neben den Versuchsflächen. Die geschätzten Ertragsänderungen für die verschiedenen Flächengrößen unter Einbezug von Ernteverlusten durch gefällte Ölpalmen sowie Effekte auf benachbarte Ölpalmen deuten darauf hin, dass die erhöhten Ernteerträge insbesondere in großen Bauminseln mindestens das Fällen von Ölpalmen kompensiert haben. Diese Ergebnisse, die in der frühen Phase der Bauminseletablierung erzielt wurden, sind vielversprechend für die Erarbeitung nachhaltiger Managementoptionen für Ölpalmplantagen, die ökologische und ökonomische Funktionen in Einklang bringen. Die anfänglichen Auswirkungen waren stärker und insbesondere aus ökonomischer Perspektive profitabler als ich erwartet habe. Die Nachbarschaftseffekte und die Auswirkungen der experimentell veränderten Variablen waren bislang jedoch überwiegend schwach. Ich erwarte, dass diese Auswirkungen mit der Zeit stärker ausgeprägt sein werden. Durch Erkenntnisse, die aus zukünftigen Langzeitbeobachtungen des Experiments, das ich in dieser Doktorarbeit vorstelle, gewonnen werden, können Wissenslücken geschlossen werden. Somit kann die Ausarbeitung von Managementrichlinien für von Ölpalmen dominierten Landschaften ermöglicht werden, die sowohl ökologisch verbessert als auch ökonomisch lohnenswert sind. Diese Doktorarbeit stellt einen wesentlichen Beitrag zur generellen Beurteilung des Experiments dar, wodurch darüber hinaus auch neue Erkenntnisse für die Renaturierungswissenschaft gewonnen werden können.

Globally land use change (LUC) with increasing in land use intensity has led to a decrease in soil organic matter (SOM). The reduction of soil C stock across LUC has been accompanied by a destabilization of soil structure and increases the soil erosion susceptibility. The destabilized soil structure is also concomitant with a loss of soil biodiversity and in particular, soil macroinvertebrate community. The focus of this dissertation is the effect of LUC with increasing in land use intensity on soil macrofauna, aggregation and SOM allocation. We hypothesized that LUC reduces soil macrofaunal biodiversity and consequently decreases soil engineering processes, resulting in destabilization of soil structure and impairing the ability of soil to physically protect SOM from decomposition inside stable aggregates, finally leading to depleted SOC stocks across LUC. In order to test these hypotheses, we conducted a field survey in 3 chronosequences of land use comprising native vegetation (NV), pasture (PA), and sugarcane crop (SC) in Central-Southern Brazil. This land use sequence provides a gradient of land use intensity and is projected to add 6.4 Mha of new sugarcane areas in Brazil by 2021. At each sampling point soil blocks of 25 X 25 cm and 5 x 5 cm to 10 cm depth were simultaneously collected from 0-10 cm, 10-20 cm, and 20-30 cm soil layers, for macrofauna isolation and aggregate fractionation, respectively. Within a radius of 6 meters around each sampling point, 12 subsamples were also collected from the same soil layers, and combined for total soil C and N concentration. An average reduction of 89% in the density of the soil macrofaunal community was observed when SC replaces PA, and a loss of 39% in the diversity of macrofauna groups. Our findings showed that, over a range of soil textures (16 - 66% clay), such biodiversity loss was strongly correlated with the destabilization of soil structure across LUC, whereas soil texture was not so. These observations consistently indicate that the abundance of detritivore soil animals, especially earthworms and termites, may be a significant predictor of soil structure transformations across LUC in tropical environments. Moreover, the sharp reduction in the abundance of earthworms was strongly positively correlated with the decrease in intra macroaggregate-associated C. As a result, after more than 20 years of sugarcane crop there were losses of 40 and 35% of C and N stocks, respectively, resulting in a rate of C emission of 1.3 Mg ha-1 yr-1. This C loss primarily occurred in the macroaggregate-associated C, as a result of the faster macroaggregate turnover under SC. In summary, the results presented here provide a mechanistic explanation as to why there is soil C depletion when increasing land use intensity in tropical soils: the huge reduction in the abundance of soil engineering invertebrates after LUC impair the ability of soil to physically protect SOM from decomposition inside stable aggregates, and therefore it is a primary mechanism controlling the soil C stocks\' depletion across LUC. / Globalmente mudanças no uso da terra (MUT) com o aumento da intensidade de uso do solo têm levado a uma diminuição da matéria orgânica do solo (MOS). A redução do estoque de C do solo após MUT tem sido acompanhada por uma desestabilização da estrutura do solo e aumento da susceptibilidade a erosão. A desestabilização da estrutura também é concomitante com a perda da biodiversidade do solo e, em particular, da comunidade de macroinvertebrados do solo. O foco deste trabalho é o efeito de MUT com aumento na intensidade de uso do solo sobre a macrofauna do solo, agregação e alocação da MOS. Nossa hipótese é que MUT reduz a diversidade da macrofauna do solo e, conseqüentemente, diminui os processos de engenharia de solo, resultando na desestabilização da estrutura do solo e prejudicando a capacidade do solo para proteger fisicamente a MOS da decomposição dentro de agregados estáveis, finalmente levando a redução dos estoques de C após MUT. Foi realizada uma pesquisa em 3 cronosseqüências de uso da terra que compreendem vegetação nativa (NV), pastagem (PA), e cana-de-açúcar (CA) na região Centro-Sul do Brasil. Esta MUT fornece um gradiente de intensidade de uso do solo e é projetada para adicionar 6,4 Mha de novas áreas de CA no Brasil até 2021. Em cada ponto de amostragem de solo blocos de 25 x 25 cm e 5 x 5 cm a 10 cm de profundidade foram coletados simultaneamente das camadas 0-10 cm, 10-20 cm e 20-30 cm de solo, para isolamento da macrofauna e fracionamento de agregados, respectivamente. Foi observada uma redução média de 89% na densidade da comunidade da macrofauna quando CA substitui PA, e uma perda de 39% da diversidade de grupos. Nossos resultados mostraram que, em um intervalo de texturas do solo (16-66% de argila), tal perda de biodiversidade foi fortemente correlacionada com a desestabilização da estrutura do solo após MUT. Estas observações indicam consistentemente que a abundância de animais detritívoros, especialmente minhocas e cupins, pode ser um preditor significativo de transformações da estrutura do solo em MUT. Além disso, a forte redução na abundância de minhocas foi fortemente e positivamente correlacionada com a diminuição do C alocado intra macroagregados. Como resultado, após mais de 20 anos de cultura de CA houve perdas de 40 e 35% dos estoques de C e N, respectivamente, resultando em uma taxa de emissão de C de 1,3 Mg ha-1 ano-1. Esta perda de C ocorreu principalmente no C associado aos macroagregados, como um resultado da reciclagem mais rápida dos macroagregados sob CA. Em resumo, os resultados aqui apresentados fornecem uma explicação mecanicista a respeito de porque há esgotamento do C do solo quando aumenta-se a intensidade de uso do solo em ambientes tropicais: a enorme redução na abundância de invertebrados \"engenheiros do solo\" após MUT prejudica a capacidade do solo para proteger fisicamente a MOS da decomposição dentro de agregados estáveis, e, portanto, é um mecanismo primário controlando a redução dos estoques de C no solo relacionada a MUT.

Tropical forests restoration is an important tool for climate change mitigation and biodiversity conservation. We can ally both of these elements, according to the biodiversity and ecosystem (BEF) functioning theory, which says that diversity enhances ecosystem functions, as primary productivity. Nevertheless, the greatest part of BEF studies up to very recently have focused on grasslands and not on as complex ecosystems as tropical forests. It is necessary to better understand above- and below-ground processes through which biodiversity acts on ecosystem functions. This work aimed to investigate effects of tree richness on both above- and below-ground ecological processes. It was based on two tropical forests undergoing restoration, in Sardinilla (Panama) and in Anhembi (Brazil). The former was especially designed for BEF studies and allowed to untangle effects of biodversity on ecosystem functions. The latter had more than a hundred species in plots and permitted investigation of the effects of high tree richness levels. In both Sardinilla and Anhembi, we investigated if tree richness levels affected an above-ground ecological process, light interception, and which mechanisms could be related to it. Richness could enhance light interception and mechanisms as spatial (horizontal and vertical) and temporal light distribution. It promoted both selection and complementarity effects. In Anhembi, we investigated if species richness influenced below-ground processes related to soil carbon stocks. Stand richness enhanced fine root production and stock. Effects of stand number of species on litter decomposition and stock were not linear. Richness of litter content, however, did not affect its decomposition rates. Number of stand species did not influence litter production. Differences of litter production, stock and fine root production among distinct richness levels did not change over the time. However, distribution of fine roots over the space, within different layers of soil, was affected by number of tree species. We concluded that even very high richness levels could not saturate some of the ecological processes studied. Diversity acted on both above- and below-ground processes, in various and sometimes opposite ways, counting on multi-direction feedbacks. It is very important to understand these mechanisms in order to potencialize biodiversity convervation and carbon sequestration by tropical forest restoration. Future studies may focus on untangling effects of diversity on below-ground processes (which have not been exhaustively explored in research), on understanding how high diversity levels affects natural regeneration and on investigationg functional traits provided by different species. / A restauração de florestas tropicais é uma importante ferramenta para a mitigação de mudanças climáticas e conservação de biodiversidade. Essas duas medidas podem ser aliadas, de acordo com a teoria de biodiversidade e funcionamento de ecossistemas (BEF, do inglês: biodiversity and ecosystem functioning), segundo a qual a diversidade pode favorecer funções do ecossistema, como a produtividade primária. Entretanto, a maior parte dos estudos de BEF até muito recentemente focaram em campos de gramíneas e não em ecossistemas tão complexos quanto florestas tropicais. É necessário entender tanto processos acima quanto abaixo do solo pelos quais a biodiversidade atua no funcionamento de ecossistemas. Este trabalho teve como objetivo verificar o efeito da riqueza de espécies arbóreas em processes ecológicos acima e abaixo do solo. Ele se baseou em duas áreas de estudo, em Sardinilla (Panamá) e em Anhembi (Brasil). A primeira foi especialmente projetada para estudos de BEF e permitiu destrinchar efeitos da biodiversidade em funções do ecossistema. A segunda possuía parcelas com mais de cem espécies, permitindo explorar os efeitos de altos níveis de riqueza. Tanto em Sardinilla quanto em Anhembi, investigamos se a riqueza de espécies arbóreas influenciou um processo ecológico acima do solo, a interceptação de luz, bem como mecanismos que podem estar associados a ele. A riqueza de espécies aumentou a interceptação de luz pelo dossel e estimulou mecanismos como a distribuição de luz ao longo do espaço (horizontal e vertical) e tempo. Ela promoveu tanto efeito de seleção quanto de complementaridade. Na área de Anhembi, investigamos se a riqueza de espécies influenciou processes abaixo do solo relacionados ao estoque de carbono nesse compartimento. A riqueza no dossel aumentou a produção e o estoque de raízes finas. Número de espécies do dossel teve efeito não linear sobre taxas de decomposição e estoque de serapilheira. A riqueza do conteúdo da serapilheira, contudo, não influenciou sua decomposição. O número de espécies do dossel também não influenciou a produção de serapilheira. As diferenças de produção e estoque de serapilheira e de produção de raízes finas, entre diferentes níveis de riquezas, não se alteraram ao longo do tempo. Contudo, o número de espécies arbóreas promoveu maior distribuição de raízes finas em diferentes camadas do solo. Concluímos que elevados níveis de riqueza não saturaram alguns processes ecológicos estudados. A diversidade foi capaz de atuar em processos tanto acima quanto abaixo do solo, por vários meios, muitas vezes em sentidos opostos, contando com feedbacks multidirecionais. É muito importante entender esses mecanismos para potencializar a conservação da biodiversidade e a provisão de funções ecossistêmicas, no processo de restauração de florestas tropicais, em um contexto internacional de necessidade de mitigação de mudanças climáticas. Estudos futuros devem focar em efeitos da diversidade em processos abaixo do solo (que são os menos abordados em estudos até o momento), em entender como altos níveis de diversidade podem afetar a regeneração natural em florestas e em explorar os atributos funcionais apresentados por cada espécie.

O funcionamento dos ecossistemas é o fluxo de matéria e de energia ao longo dos seus componentes bióticos e abióticos. A manutenção de tal funcionamento é crucial para promover os serviços ecossistêmicos dos quais a humanidade depende. Atividades antrópicas, como a agricultura e a urbanização podem alterá-los significativamente e, por isso, o entendimento dos impactos positivos e negativos de tais alterações bem como o desenvolvimento de ferramentas de avaliação da biodiversidade são importantes para a conservação. A diversidade funcional é um componente da diversidade biológica que leva em conta o papel que cada espécie desempenha no ecossistema e, portanto, deve refletir o funcionamento do ecossistema de forma mais acurada. Esta tese teve por objetivos relacionar o funcionamento do ecossistema e os seus serviços a ações antrópicas, bem como propor a abordagem funcional como indicadora da biodiversidade de áreas naturais. No primeiro capítulo, avalio padrões espaciais e temporais da exposição dos tipos de vegetação que ocorrem no Brasil à luz artificial, um dos principais símbolos da urbanização e da vida moderna. Os resultados mostram que a maioria deles apresenta alguma porcentagem de sua área expostos à luz artificial. Em alguns deles, porém, ainda é possível encontrar o brilho natural do céu noturno, o que nos permite sugerir a elaboração de políticas de desenvolvimento de distribuição de luz com foco no mínimo impacto, ao contrário das políticas de mitigação adotadas por países onde a iluminação é excessivamente difundida. No segundo capítulo mapeio padrões espaciais e temporais da provisão de dois serviços ecossistêmicos (polinização e estoque de carbono) e um serviço ambiental (espaço para viver) e avalio os impactos da agricultura sobre estes serviços, na área coberta pelo cerrado, na região central do Brasil. Os serviços ecossistêmicos avaliados descaíram significativamente ao longo do tempo e a região conhecida como MATOPIBA (entre os estados do Maranhão, Tocantins, Piauí e Bahia) foi identificada como um importante remanescente. Por isso, recomendo a criação de unidades de conservação de proteção integral na área. Identifiquei também sobreposição da disponibilidade destes serviços com áreas indígenas, o que ressalta a importância da manutenção destas áreas para a disponibilidade dos serviços. No terceiro capítulo exploro, por meio de revisão bibliográfica, o status da diversidade funcional na literatura científica no campo da Conservação Biológica. Os resultados permitiram visualizar o paradigma da transferência de conhecimento e identificar o potencial desta abordagem para elaboração de indicadores da biodiversidade. Por fim, no quarto capítulo, estabeleço uma lista de espécies de plantas que podem ser indicadoras da diversidade funcional em áreas cobertas por cerradão no estado de São Paulo. Tais indicadores devem ser usados em conjunto para monitoramento e diagnóstico da biodiversidade quando o objetivo é manter o funcionamento do ecossistema. As conclusões do estudo contribuem para elucidar a interferência das atividades antrópicas nos ecossistemas naturais, para levantar formas de minimizá-los e para aprimorar as formas de avaliação da biodiversidade. / Ecosystem functioning is the flow of energy and matter through the biotic and abiotic ecosystems components. The maintenance of this functioning is essential to the ecosystem services provisioning upon which humans depend. Anthropogenic activities such as agriculture and urbanisation may change it. Thus, conservation strategies rely on the understanding of the positives and negatives impacts from this changes and on the development of measurements of biodiversity. Functional diversity is a biodiversity component which considers the role of each species in the ecosystem, and, as a consequence, may reflect the ecosystem functioning more accurately. This thesis aims the establishment of the relationships between ecosystem functioning, ecosystem services and anthrogenic activities as well as purpose functional approach as biodiversity indicator of natural areas. In the first chapter, I assessed spatial and temporal patterns of exposition of the Brazilian vegetation types to the artificial light which is one of the urbanisation symbols. The results show that most of them are affected by artificial light. However, in some of them it is still possible to find a natural sky background, which allow suggesting the formulation of light distribution policies focused on minimal impact instead of mitigation, as adopted by countries where light is excessively widespread. In the second chapter, I mapped spatial and temporal patterns of two ecosystem services provisioning (pollination and carbon stocks) and one environmental service (living space) and assessed the impacts of agriculture in these services in Cerrado area, in the central region of Brazil. The ecosystem services I assessed declined significatively over time and the MATOPIBA region (around Maranhão, Tocantins, Piaui, and Bahia States) was identified as an important remint. Thus I recommended the creation of strictly protected areas in that region. I also identify overlap of these services with indigenous land, highlighting its importance for the ecosystem services provisioning. In the third chapter I explored through literature survey, the status of functional diversity in the scientific literature related to the Biological Conservation field. The results allow visualise the paradigm of knowledge transfer and identify the potential use of this approach for developing indicators of biodiversity. Finally, in the fourth chapter, we established a list of plant species that are functional diversity indicators in cerradão areas in São Paulo State. These indicators should be use as a set for the monitoring and diagnosis of biodiversity when the goal is to maintaining ecosystem functioning. The conclusions of the study contribute to clarify some lack of knowledge concerned to the impacts of human activities on ecosystem functioning, to raise means to minimise them, and to improve means of ecological integrity assessment.