Die Ökosysteme unserer Erde sind durch das rasante Artensterben infolge von Umweltveränderungen durch den Menschen und des globalen Klimawandels stark betroffen. Mit den Auswirkungen dieses Artenverlustes und der damit einhergehenden Veränderung der Diversität beschäftigt sich die heutige Biodiversitätsforschung. Spezieller wird der Effekt der Diversität auf Ökosystemprozesse wie beispielsweise den Biomasseaufbau von Primärproduzenten oder der Resistenz einer Gemeinschaft gegen die Einwanderung neuer Arten untersucht. Die Quantifizierung des Einflusses der Diversität auf die Primärproduktion und das Verständnis der zugrunde liegenden Mechanismen ist von besonderer Wichtigkeit. In terrestrischen Pflanzengemeinschaften wurde bereits ein positiver Diversitätseffekt auf die Gemeinschaftsbiomasse beobachtet. Dies wird hauptsächlich durch den Komplementaritäts- und/oder den Dominanzeffekt erklärt. Die Komplementarität zwischen Arten ist beispielsweise bei Unterschieden in der Ressourcenausnutzung gegeben (z.B. unterschiedliche Wurzeltiefen). Diese kann zu einer besseren Nährstoffausnutzung in diverseren Gemeinschaften führen, die letztlich deren höhere Biomassen erklärt. Der Dominanzeffekt hingegen beruht auf der in diverseren Gemeinschaften höheren Wahrscheinlichkeit, eine hochproduktive Art anzutreffen, was letztlich die höhere Biomasse der Gemeinschaft verursacht. Diversitätseffekte auf Ökosystemprozesse wurden bisher hauptsächlich auf der Gemeinschaftsebene untersucht. Analysen über die Reaktionen, die alle Arten einer Gemeinschaft einschließen, fehlen bisher. Daher wurde der Einfluss der Diversität auf die individuelle Performance von Pflanzenarten innerhalb des Biodiversitätsprojektes „Das Jena Experiment“ untersucht. Dieses Experiment umfasst 60 Arten, die charakteristisch für Mitteleuropäische Graslandschaften sind. Die Arten wurden in die 4 funktionellen Gruppen Gräser, kleine Kräuter, große Kräuter und Leguminosen eingeteilt. Im Freilandversuch zeigte sich, dass mit steigender Artenzahl die individuelle Pflanzenhöhe zunahm, während die individuelle oberirdische Biomasse sank. Der positive Diversitätseffekt auf die pflanzliche Gemeinschaftsbiomasse kann folglich nicht auf der individuellen oberirdischen Biomassezunahme beruhen. Überdies reagierten die einzelnen funktionellen Gruppen und sogar die einzelnen Arten innerhalb einer funktionellen Gruppe unterschiedlich auf Diversitätsveränderungen. Folglich ist zu vermuten, dass einige Ökosystemprozesse auf Gemeinschaftsebene durch die Reaktionen von bestimmten funktionellen Gruppen bzw. Arten hervorgerufen werden. Diversitätseffekte auf Gemeinschaftsbiomassen wurden bislang hauptsächlich mit terrestrischen Pflanzen und weniger mit frei-schwebenden Algenarten (Phytoplankton) erforscht. Demzufolge wurde der Einfluss der Diversität auf die Biomasse von Phytoplankton-Gemeinschaften experimentell untersucht, wobei es sowohl zu negativen als auch positiven Diversitätseffekten kam. Eine negative Beziehung zwischen Diversität und Gemeinschaftsbiomasse zeigte sich, wenn schnell-wüchsige Algenarten nur geringe Biomassen in Mono- und Mischkultur aufbauten. Die vorhandenen Nährstoffe in der Mischkultur wurden von den schnell-wüchsigen Arten monopolisiert und folglich standen sie den langsam-wüchsigen Algenarten, welche viel Biomasse in Monokultur aufbauten, nicht mehr zur Verfügung. Zu einem positiven Diversitätseffekt auf die Gemeinschaftsbiomasse kam es, wenn die Artengemeinschaft eine positive Beziehung zwischen Wachstumsrate und Biomasse in Monokultur zeigte, sodass die schnell-wüchsige Algenarten viel Biomasse aufbauten. Da diese schnell-wüchsigen Algen in der Mischkultur dominant wurden, bestand die Gemeinschaft letztlich aus hoch-produktiven Algenarten, was zu einer erhöhten Gesamtbiomasse führte. Diese beiden Versuchsansätze verdeutlichen Mechanismen für die unterschiedlichen Reaktionen der Gemeinschaften auf Diversitätsveränderungen, welche auch für terrestrische Pflanzengemeinschaften gefunden wurden. Ein anderer wichtiger Ökosystemprozess, der von der Diversität beeinflusst wird, ist die Anfälligkeit von Gemeinschaften gegenüber invasiven Arten (Invasibilität). Die Invasibilität wird von einer Vielzahl von Faktoren beeinflusst und demzufolge wurde der Effekt der Diversität und der Produktivität (Nährstoffgehalt) auf die Invasibilität von Phytoplankton-Gemeinschaften in An- und Abwesenheit eines Herbivoren untersucht. Die zwei funktionell unterschiedlichen invasiven Arten waren die Blaualge Cylindrospermopsis raciborskii (schlecht fressbar) und der Phytoflagellat Cryptomonas sp. (gut fressbar). Es zeigte sich, dass der Fraßdruck, welcher selber durch die Produktivität beeinflusst wurde, einen bedeutenden Effekt auf die Invasibilität von Phytoplankton-Gemeinschaften hat. Die funktionellen Eigenschaften der invasiven und residenten Arten waren zudem bedeutender als die Artenzahl. / To date, positive relationships between diversity and community biomass have been mainly found, especially in terrestrial ecosystems due to the complementarity and/or dominance effect. In this thesis, the effect of diversity on the performance of terrestrial plant and phytoplankton communities was investigated to get a better understanding of the underlying mechanisms in the biodiversity-ecosystem functioning context. In a large grassland biodiversity experiment, the Jena Experiment, the effect of community diversity on the individual plant performance was investigated for all species. The species pool consisted of 60 plant species belonging to 4 functional groups (grasses, small herbs, tall herbs, legumes). The experiment included 82 large plots which differed in species richness (1-60), functional richness (1-4), and community composition. Individual plant height increased with increasing species richness suggesting stronger competition for light in more diverse communities. The aboveground biomass of the individual plants decreased with increasing species richness indicating stronger competition in more species-rich communities. Moreover, in more species-rich communities plant individuals were less likely to flower out and had fewer inflorescences which may be resulting from a trade-off between resource allocation to vegetative height growth and to reproduction. Responses to changing species richness differed strongly between functional groups and between species of similar functional groups. To conclude, individual plant performance can largely depend on the diversity of the surrounding community. Positive diversity effects on biomass have been mainly found for substrate-bound plant communities. Therefore, the effect of diversity on the community biomass of phytoplankton was studied using microcosms. The communities consisted of 8 algal species belonging to 4 functional groups (green algae, diatoms, cyanobacteria, phytoflagellates) and were grown at different functional richness levels (1-4). Functional richness and community biomass were negatively correlated and all community biomasses were lower than their average monoculture biomasses of the component species, revealing community underyielding. This was mainly caused by the dominance of a fast-growing species which built up low biomasses in monoculture and mixture. A trade-off between biomass and growth rate in monoculture was found for all species, and thus fast-growing species built up low biomasses and slow-growing species reached high biomasses in monoculture. As the fast-growing, low-productive species monopolised nutrients in the mixtures, they became the dominant species resulting in the observed community underyielding. These findings suggest community overyielding when biomasses of the component species are positively correlated with their growth rates in monocultures. Aquatic microcosm experiments with an extensive design were performed to get a broad range of community responses. The phytoplankton communities differed in species diversity (1, 2, 4, 8, and 12), functional diversity (1, 2, 3, and 4) and community composition. The species/functional diversity positively affected community biomass, revealing overyielding in most of the communities. This was mainly caused by a positive complementarity effect which can be attributed to resource use complementarity and/or facilitative interaction among the species. Overyielding of more diverse communities occurred when the biomass of the component species was correlated positively with their growth rates in monoculture and thus, fast-growing and high-productive species were dominant in mixtures. This and the study mentioned above generated an emergent pattern for community overyielding and underyielding from the relationship between biomass and growth rate in monoculture as long as the initial community structure prevailed. Invasive species can largely affect ecosystem processes, whereas invasion is also influenced by diversity. To date, studies revealed negative and positive diversity effects on the invasibility (susceptibility of a community to the invasion by new species). The effect of productivity (nutrient concentration ranging from 10 to 640 µg P L-1), herbivory (presence/absence of the generalist feeder) and diversity (3, 4, 6 species were randomly chosen from the resident species pool) on the invasibility of phytoplankton communities consisting of 10 resident species was investigated using semi-continuous microcosms. Two functionally diverse invaders were chosen: the filamentous and less-edible cynaobacterium C. raciborskii and the unicellular and well-edible phytoflagellate Cryptomonas sp. The phytoflagellate indirectly benefited from grazing pressure of herbivores whereas C. raciborskii suffered more from it. Diversity did not affect the invasibility of the phytoplankton communities. Rather, it was strongly influenced by the functional traits of the resident and invasive species.

Human-induced alterations of the environment are causing biotic changes worldwide, including the extinction of species and a mixing of once disparate floras and faunas. One type of biological communities that is expected to be particularly affected by environmental alterations are herb layer plant communities of fragmented forests such as those in the west European lowlands. However, our knowledge about current changes in species diversity and composition in these communities is limited due to a lack of adequate long-term studies. In this thesis, I resurveyed the herb layer communities of ancient forest patches in the Weser-Elbe region (NW Germany) after two decades using 175 semi-permanent plots. The general objectives were (i) to quantify changes in plant species diversity considering also between-community (β) and functional diversity, (ii) to determine shifts in species composition in terms of species’ niche breadth and functional traits and (iii) to find indications on the most likely environmental drivers for the observed changes. These objectives were pursued with four independent research papers (Chapters 1-4) whose results were brought together in a General Discussion. Alpha diversity (species richness) increased by almost four species on average, whereas β diversity tended to decrease (Chapter 1). The latter is interpreted as a beginning floristic homogenization. The observed changes were primarily the result of a spread of native habitat generalists that are able to tolerate broad pH and moisture ranges. The changes in α and β diversity were only significant when species abundances were neglected (Chapters 1 and 2), demonstrating that the diversity changes resulted mainly from gains and losses of low-abundance species. This study is one of the first studies in temperate Europe that demonstrates floristic homogenization of forest plant communities at a larger than local scale. The diversity changes found at the taxonomic level did not result in similar changes at the functional level (Chapter 2). The likely reason is that these communities are functionally “buffered”. Single communities involve most of the functional diversity of the regional pool, i.e., they are already functionally rich, while they are functionally redundant among each other, i.e., they are already homogeneous. Independent of taxonomic homogenization, the abundance of 30 species decreased significantly (Chapter 4). These species included 12 ancient forest species (i.e., species closely tied to forest patches with a habitat continuity > 200 years) and seven species listed on the Red List of endangered plant species in NW Germany. If these decreases continue over the next decades, local extinctions may result. This biotic impoverishment would seriously conflict with regional conservation goals. Community assembly mechanisms changed at the local level particularly at sites that experienced disturbance by forest management activities between the sampling periods (Chapter 3). Disturbance altered community assembly mechanisms in two ways: (i) it relaxed environmental filters and allowed the coexistence of different reproduction strategies, as reflected by a higher diversity of reproductive traits at the time of the resurvey, and (ii) it enhanced light availability and tightened competitive filters. These limited the functional diversity with respect to canopy height and selected for taller species. Thirty-one winner and 30 loser species, which had significantly increased or decreased in abundance, respectively, were characterized by various functional traits and ecological performances to find indications on the most likely environmental drivers for the observed floristic changes (Chapter 4). Winner species had higher seed longevity, flowered later in the season and had more often an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, to be more susceptible to deer browsing and to have a performance optimum at higher soil pH values compared to winner species. Multiple logistic regression analyses indicated that disturbances due to forest management interventions were the primary cause of the species shifts. As one of the first European resurvey studies, this study provides indications that an enhanced browsing pressure due to increased deer densities and increasingly warmer winters are important drivers. The study failed to demonstrate that eutrophication and acidification due to atmospheric deposition substantially drive herb layer changes. The restriction of the sample to the most base-rich sites in the region is discussed as a likely reason. Furthermore, the decline of several ancient forest species is discussed as an indication that the forest patches are still paying off their “extinction debt”, i.e., exhibit a delayed response to forest fragmentation. / Umweltveränderungen beeinträchtigen weltweit die Artenvielfalt. Die Lebensgemeinschaften fragmentierter Lebensräume gelten als besonders anfällig für Veränderungen. In dieser Arbeit wurden Untersuchungen an Krautschichtgemeinschaften historisch alter Waldfragmente im Elbe-Weser-Dreieck nach zwei Jahrzehnten wiederholt. Ziel war es anhand von 175 semi-permanenten Aufnahmeflächen (i) die Veränderungen der Pflanzenartendiversität zu quantifizieren, (ii) Artenverschiebungen in Bezug auf Nischenbreite und funktionale Merkmale festzustellen und (iii) Hinweise auf die verantwortlichen Umweltveränderungen zu finden. Die α-Diversität (Artenzahl) stieg durchschnittlich um vier Arten an. Die β-Diversität (Artenturnover zwischen den Flächen) nahm tendenziell ab. Letzteres wird als Beginn einer floristischen Homogenisierung interpretiert. Diese Studie ist eine der ersten im gemäßigten Europa, die eine floristische Homogenisierung von Waldpflanzengemeinschaften auf einer größeren als der lokalen Ebene aufzeigt. Die Diversitätsveränderungen auf taxonomischer Ebene führten nicht zu ähnlichen Veränderungen auf funktionaler Ebene. Bereits einzelne Gemeinschaften wiesen den Großteil der funktionalen Vielfalt des regionalen Artenpools, also ein Maximum an funktionaler Diversität auf. Gleichzeitig waren sie untereinander funktional redundant, also bereits homogen. Die mit der beginnenden taxonomischen Homogenisierung verbundene floristische Verarmung wird als gering eingestuft, da die Homogenisierung primär das Ergebnis der Zuwanderung häufig vorkommender Standortgeneralisten war. Unabhängig von der Homogenisierung gingen 30 Arten signifikant in ihrer Abundanz zurück, darunter 12 an historisch alte Wälder gebundene Arten sowie sieben Rote-Liste-Arten. Ein weiterer Rückgang oder ein lokales Aussterben dieser Arten stünde im Widerspruch zu regionalen Naturschutzzielen. Nullmodelltests und der Vergleich funktionaler und taxonomischer Diversitätskomponenten lassen auf regionaler Ebene auf eine zeitliche Konstanz der grundlegenden Mechanismen der Artenvergesellschaftung schließen. Auf der lokalen Ebene veränderten sich die Vergesellschaftungsmechanismen erheblich, insbesondere auf forstwirtschaftlich gestörten Standorten. Einerseits ermöglichte dort eine Abschwächung der Umweltfilter die Koexistenz von Arten mit unterschiedlichen Reproduktionsstrategien. Andererseits führte die erhöhte Lichtverfügbarkeit zu einer Verstärkung der Konkurrenzfilter und einer Selektion hochwüchsiger Arten. Gewinner- und Verliererarten wurden anhand funktionaler Merkmale und ihres ökologischen Verhaltens charakterisiert, um Hinweise auf die verantwortlichen Umweltveränderungen zu finden. Gewinnerarten wiesen eine höhere Langlebigkeit der Samen auf, blühten später in der Vegetationsperiode und hatten öfter eine ozeanische Verbreitung. Verliererarten hatten eine höhere spezifische Blattfläche, einen höheren Attraktivitätswert als Wildäsung und ein ökologisches Optimum bei höheren pH-Werten. Logistische Regressionsanalysen zeigen, dass Störung durch forstwirtschaftliche Eingriffe hauptverantwortlich für die Artenverschiebungen war. Zusätzlich liefert diese Wiederholungsstudie als eine der ersten in Europa Hinweise darauf, dass ein erhöhter Äsungsdruck sowie zunehmend mildere Winter entscheidende Einflussfaktoren darstellen. Der Rückgang mehrerer an historisch alte Wälder gebundener Arten wird als Anzeichen für eine verspätete Reaktion auf die Waldfragmentierung diskutiert.

Grünland spielt eine große Rolle in der Landnutzung und nimmt ein Drittel der landwirtschaftlich genutzten Fläche von Europa ein. Als Konsequenz der Intensivierung landwirtschaftlicher Bewirtschaftungsmaßnahmen der letzten 60 Jahre nahm die Produktivität des Grünlands zu während die Diversität dieser Systeme abnahm. In Grünland-Ökosystemen spielt Stickstoff (N) eine Schlüsselrolle – N bedingt die Primärproduktion und beeinflusst die Biodiversität. Zudem kann eine steigende N-Verfügbarkeit gasförmige Emissionen, wie z.B. Distickstoffoxid (N2O) und Stickstoffmonoxid (NO) fördern, die eine große Rolle in der Atmosphäre spielen und zur globalen Erwärmung beitragen. Eine höhere Nitratauswaschung (NO3-) aus Böden kann eine Gefahr für die Grundwasserqualität sein. N-Verluste durch Ausgasung von N2O und NO sowie NO3--Auswaschung sind dabei die Folgen der mikrobiellen Prozesse Denitrifikation und Nitrifikation. In dieser Studie haben wir den Effekt von unterschiedlichen Bewirtschaftungsintensitäten und funktioneller Pflanzendiversität auf die N-Verluste und Ökosystemfunktionen untersucht. Die Studie ist Teil des Excellenzclusters „Funktionelle Biodiversitätsforschung“ der Georg-August-Universität Göttingen und wurde durch das Niedersächsische Ministerium für Wissenschaft und Kultur finanziert. Die Studie wurde im Rahmen von zwei interdisziplinären Projekten (BIOMIX & GRASSMAN) von 2008 bis 2010 im Solling, Niedersachsen, Deutschland durchgeführt. Wir untersuchten von Rindern und Schafen beweidetes Grünland (BIOMIX) und gemähtes Grünland mit unterschiedlichen Bewirtschaftungsintensitäten (GRASSMAN). In beiden Projekten wurde die funktionelle Pflanzendiversität durch Herbizide eingestellt. Der Fokus unserer Arbeit lag auf den N-Verlusten (N2O and NO Emissionen, NO3--Auswaschung) und der N Dynamik (Netto und Brutto Mineralisation). In GRASSMAN berechneten wir zusätzlich die N-Nutzungseffizienz und die N-Rückhalteeffizienz auf Ökosystemebene. Dabei ist die N-Nutzungseffizienz das Produkt der Aufnahmeeffizienz (definiert als N-Aufnahme der Pflanze pro verfügbares N) und der N-Nutzungseffizienz auf Pflanzenebene (definiert als Produktivität pro N-Aufnahme der Pflanze). Darüber hinaus berechnen wir N-Rückhalteeffizienz in Böden als einen Index, der das Verhältnis von N-Verlusten zu dem im Grünland verbleibenden N beschreibt. In BIOMIX haben wir die Auswirkung von Beweidung und Pflanzenarten-zusammensetzung auf N2O and NO Emissionen untersucht. Die mit einem Herbizid gegen Dikotyle vorbehandelten Weiden wurden mit Rindern oder Schafen Rotationsweise beweidet. Mittlere N2O Emissionen lagen bei 38.7 µg N2O-N m-2 h-1, mittlere NO Emissionen betrugen 2.4 µg NO-N m-2 h-1. Kumulative NO-N Emissionen waren höher auf den von Schafen beweideten Flächen als auf den von Rindern beweideten Flächen. In einem kontrollierten Applikations-Experiment führte die Behandlung mit Rinderurin zu höheren N2O Emissionen als die Behandlung mit Schafurin. Die Emissionshöchstwerte von 1921 µg N2O-N m-2 h-1 bei Behandlung mit Rinderurin im Vergleich zu 556 µg N2O-N m-2 h-1 bei Schafurin standen im Zusammenhang mit unterschiedlichen N-Einträgen pro Ausscheidung der Tiere. Die Emissionshöchstwerte der mit Dung behandelten Flächen waren im Vergleich mit den jeweiligen Urinbehandlungen viel geringer. Die N2O Emissionsfaktoren betrugen 0.4% für Rinderurin, 0.5% für Schafurin, 0.05% für Rinderdung und 0.09% für Schafdung. Sowohl das Beweidungs-Experiment, als auch das kontrollierte Applikations-Experiment zeigten, dass die Pflanzenartenzusammensetzung auf N-Emissionen im Vergleich zum Einfluss der Weidetierart auf N-Emissionen unbedeutend war. Trotz höherer N-Einträge auf Rinderweiden waren die N-Emissionen aus der Schafbeweidung höher. Wir führten dies auf die gleichmäßigere Verteilung von Schafs-Exkrementen im Vergleich zu Rindern-Exkrementen zurück. In GRASSMAN untersuchten wir die Auswirkungen von unterschiedlichen Bewirtschaftungsregimen (Düngung und Schnittintensität) und Pflanzenarten-zusammensetzung auf die N-Verluste (N2O Emissionen, NO3- Auswaschung) und die N-Dynamik (Netto und Brutto Mineralisation) und kalkulierten die N-Nutzungseffizienz und die N-Rückhalteeffizienz. Ein dreifaktorielles Design mit folgenden Faktoren wurde über einen Zeitraum von zwei Jahren etabliert: Düngung (180 – 30 – 100 kg NPK ha-1 yr-1 und keine Düngung), Schnittintensität (ein- und dreimal pro Jahr) und Pflanzenartenzusammensetzung (eine unbehandelte Kontrolle, eine Dikotyl-erhöhte und eine Monokotyl-erhöhte Grasnarbe). In 2009 wurden die N2O Emissionen erheblich von beiden Bewirtschaftungsfaktoren (Düngung und Schnittintensität) beeinflusst, während in 2010 nur die Düngung die N2O Emissionen beeinflusste. In 2009 wurden NO3- Auswaschungsverluste durch Düngung und in 2010 von beiden Bewirtschaftungsfaktoren (Düngung und Schnittintensität) beeinflusst. Die Netto N-Mineralisation Raten wurden in 2009 nur von der Düngung beeinflusst. In 2010, zeigte nicht nur die Düngung, sondern auch die Schnittintensität einen Einfluss auf die Netto N-Mineralisation Raten. Weder die Bewirtschaftung (Düngung) noch die Pflanzenartenzusammensetzung hatte einen Einfluss auf die Brutto N-Mineralisation. Die N-Nutzungseffizienz wurde vor allem durch die Düngung und als weiterer Faktor durch die Schnittintensität in 2009 beeinflusst, welche 41% bzw. 3% der Varianz erklärten. In 2010 hatte die Düngung mit 24% der erklärten Varianz einen geringeren Effekt auf die N-Nutzungseffizienz, während die Auswirkungen der Schnittintensität (12%) und die Pflanzenartenzusammensetzung (6%) stärker ausgeprägt waren. Die N-Nutzungseffizienz war auf ungedüngten Flächen größer als auf gedüngten, in den dreimal geschnittenen Flächen höher als in den einmal geschnittenen, und in der unbehandelten Kontrolle höher als in der Monokotyl-erhöhte oder Dikotyl-erhöhte Grasnarbe. Düngung verringert die N-Nutzungseffizienz durch die Abnahme in der N-Aufnahmeeffizienz und der N-Nutzungseffizienz auf Pflanzenebene, während die Schnittintensität und die Pflanzenartenzusammensetzung nur durch die N-Aufnahmeeffizienz beeinflusst werden. Die N-Rückhalteeffizienz wurde nur für 2010 berechnet und wurde durch die Düngung und die Pflanzenartenzusammensetzung mit 22% und 17% der erklärten Varianz beeinflusst. N-Rückhalteeffizienz nahm in der Reihenfolge unbehandelte Kontrolle > Dikotyl-erhöhte > Monokotyl-erhöhte Grasnarbe mit einem signifikanten Unterschied zwischen der unbehandelten Kontrolle und der Monokotyl-erhöhten Grasnarbe ab. Die N-Rückhalteeffizienz ist mit dem mikrobiellen Ammonium (NH4+) und der mikrobiellen Biomasse hoch und mit der N-Aufnahme der Pflanzen nur gering korreliert, was die Bedeutung der mikrobiellen N Retention im System Boden-Pflanze unterstreicht. Unsere Ergebnisse zeigen, dass die Bewirtschaftung der wichtigste und bestimmende Faktor der Ökosystemfunktionen eines Grünlands ist. Düngung, Schnittintensität und Beweidung beeinflussen die N-Nutzungseffizienz, die N-Rückhalteeffizienz und die N-Verluste. Die Zusammensetzung der botanischen Grasnarbe hat einen geringen Einfluss auf den N Kreislauf oder die N-Nutzungs- und die N-Rückhalteeffizienz. Wobei die Pflanzenartenzusammensetzung der unbehandelten Kontrolle (~70% Monokotyle und ~30% Dikotyle), die sich unter der extensiven Langzeit-Bewirtschaftung eingestellt hatte, die höchsten Effizienzen zeigte - sowohl eine Erhöhung der Monokotyledonen als auch eine Erhöhung der Dikotyledonen führte zu einer Verringerung der Effizienzen. Darüber hinaus sind N-Nutzungs- und N-Rückhalteeffizienz geeignete Werkzeuge, die sich zur Evaluierung ökologischer Nachhaltigkeit von Pflanzenartenzusammensetzungen und Management-Praktiken im Grünland eignen.

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Submitted by Helena Alves do Prado null (helenapradoeco@gmail.com) on 2017-12-08T19:34:27Z No. of bitstreams: 1 Dissertação_HAP.pdf: 5105198 bytes, checksum: 05421b9dc8b38cd4f75d69d9f2066044 (MD5) / Approved for entry into archive by Adriana Aparecida Puerta null (dripuerta@rc.unesp.br) on 2017-12-11T16:34:40Z (GMT) No. of bitstreams: 1 prado_ha_me_rcla.pdf: 5105198 bytes, checksum: 05421b9dc8b38cd4f75d69d9f2066044 (MD5) / Made available in DSpace on 2017-12-11T16:34:40Z (GMT). No. of bitstreams: 1 prado_ha_me_rcla.pdf: 5105198 bytes, checksum: 05421b9dc8b38cd4f75d69d9f2066044 (MD5) Previous issue date: 2017-07-27 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) / A importância das florestas tropicais para o ciclo de carbono global e a incerteza de suas respostas a novas condições ambientais evidenciam a necessidade de aprofundar o conhecimento acerca de como as mudanças climáticas repercutirão nas dinâmicas florestais e nos processos fisiológicos associados. Essa dissertação, vinculada ao experimento AmazonFACE, objetiva avaliar como o incremento de CO2 atmosférico poderia repercutir na produtividade e eficiência no uso da água da Floresta Amazônica; se atributos funcionais envolvidos nesses processos (vcmax e g1) seriam modificados; e se ocorreriam mudanças na configuração funcional modelada da região. Para melhor caracterizar a diversidade funcional e as diferentes estratégias de sobrevivência adotadas pelas plantas em um ecossistema hiperdiverso, desenvolveu-se o CAETÊ (Carbon and Ecosystem Functional-Trait Evaluation Model) que, diferentemente dos modelos baseados em tipos funcionais, não parametriza a priori as possíveis combinações de atributos funcionais e simula um número superior de estratégicas de vida (PLS), sendo essas amostradas semi-aleatoriamente pelo modelo. Ao avaliar a performance do CAETÊ, verificou-se que a consideração de um número elevado de PLS – em vez de uma abordagem baseada em poucos tipos funcionais – simula maiores taxas de produtividade e condutância estomática; e que esse aumento tende à estabilização, com o aumento de estratégias consideradas. A aplicação do modelo foi feita utilizando a concentração de CO2 a ser empregada no AmazonFACE, de 600 ppm. Nas simulações, verificou-se uma redução nos valores médios de vcmax e g1, assim como uma modificação nos seus padrões de distribuição. Essas alterações associaram-se a um incremento nas taxas modeladas de produtividade e no uso da água, nas florestas tropicais, próximo a 13.5% (GPP) e 14.2% (NPP); redução de 3.8% na condutância estomática; e maior eficiência no uso da água (+21.1%). O enriquecimento da atmosfera também esteve associado à redução de ~5.5% na riqueza funcional. À medida que se avança para condições ambientais sem precedentes, torna-se de grande valia o uso de novas abordagens pelos modelos vegetacionais, que possibilitem a consideração de um número superior de estratégias de sobrevivência das plantas; e a captação de possíveis alterações nas respostas de atributos funcionais essenciais e, consequentemente, na riqueza funcional dos ecossistemas. As florestas tropicais desempenham um papel essencial nessas dinâmicas, uma vez que, além de hiperdiversas, são centrais para o ciclo de carbono global. / The importance of tropical forests in the global carbon cycle and the uncertainty of the forest’s responses to new environmental conditions evidence the necessity of deepening our knowledge about how climate changes affect forest dynamics and the associated physiological processes. This dissertation, linked to the AmazonFACE experiment, aims to evaluate how increasing atmospheric levels of CO2 could influence primary productivity and water use efficiency of the Amazon Forest; if functional traits involved in such processes (vcmax e g1) would be modified; and if changes would occur in the region’s modeled functional configuration. Aiming for a better characterization of the functional diversity and the different survival strategies adopted by plants in a hyperdiverse ecosystem, we developed the CAETÊ (Carbon and Ecosystem Functional-Trait Evaluation Model) which, unlike models based on functional types, does not perform a priori parameterization of the possible combinations of functional traits and simulates a superior amount of life strategies (PLS), that are semi-randomly sampled by the model. When evaluating CAETÊ’s performance, we verified that the consideration high number of PLS – instead of a framework based in few functional types – simulates higher amounts of productivity and stomatal conductance; and that such increase tends to stabilization when more strategies are considered. The model was applied using the CO2 concentration to be used in AmazonFACE – 600ppm. Simulations showed a reduction in the average values of de vcmax and g1, as well as a change in their patterns of distribution. Such alterations are associated to an increase in modeled values of productivity and water use of tropical forests close to 13.5% (GPP) and 14.2% (NPP), a reduction of 3.8% in stomatal conductance, and higher efficiency in water use (+21.1%). The atmospheric enrichment was also associated with a ~5.5% reduction in the functional richness. As the environment advances to unprecedented conditions, it becomes clear how worthy is the use of new approaches by vegetational models that enable consideration of higher numbers of plant survival strategies; and the uptake of possible changes in responses of essential functional traits, therefore, in the functional richness of the environment. The tropical forests influence deeply such dynamics; once beyond hyperdiverse.

Muitas questões ligadas à ecologia, biologia da conservação e biogeografia estão relacionadas aos efeitos de processos naturais ou antropogênicos sobre fragmentos, manchas ou ilhas. Na atualidade, para se compreender os efeitos da fragmentação e se propor medidas efetivas de conservação, torna-se cada vez mais importante analisar os fragmentos como parte de uma matriz que inclui não somente diferentes tamanhos de remanescentes e de floresta contínua, mas também outros elementos da estrutura da paisagem. Para tanto, quando comparamos formações com diferentes características ambientais, mesmo que inseridas em regiões próximas, para entendermos os mecanismos que levam ao desaparecimento de algumas espécies e beneficiam outras, é necessário considerar níveis de organização relacionados a atributos de espécies e populações. Determinados atributos podem ser indicadores ecológicos relevantes de pressões exercidas pelo ambiente na biologia das espécies, ao mesmo tempo em que demonstram relação com a função do organismo. Essa tese, em seu primeiro capítulo avalia como questões metodológicas e analíticas podem influenciar os resultados de estudos avifaunísticos e a relação da riqueza e abundância de espécies com o tamanho de fragmentos e manchas florestais. Observamos que a relação entre a riqueza e o tamanho das áreas foi afetada pela padronização do tempo de contagem por área. A utilização de contagens com raio ilimitado fez com que o número de espécies aumentasse significativamente somente nas maiores manchas de Floresta Ombrófila Mista. Considerando-se o acréscimo de registros obtidos em cada minuto de contagem, constatou-se que com nove minutos já haviam sido obtidos grande parte dos registros de espécies e contatos de aves. No segundo capítulo, foi investigado se a riqueza rarefeita e a abundância de aves diminuem com o decréscimo da área de fragmentos de Floresta Ombrófila Densa. Paralelamente buscou-se compreender como a avifauna é influenciada por características da vegetação dos fragmentos e do entorno destes. A abundância obtida em cada fragmento esteve significativamente relacionada ao tamanho das áreas, sendo mais elevada nos fragmentos maiores. As análises relacionando a riqueza rarefeita média dos fragmentos e as variáveis ambientais selecionadas mostraram que as variáveis de mancha foram mais importantes que as variáveis relacionadas à estrutura da vegetação no interior dos fragmentos. No terceiro capítulo, avaliamos como as comunidades de aves respondem diferentemente à fragmentação em regiões originalmente florestadas e à existência de mosaicos em regiões com manchas florestais (capões), situações características das regiões onde estão inseridas, respectivamente, as Florestas Ombrófilas Densa (FOD) e Mista (FOM). As áreas de FOM e FOD foram agrupadas quando relacionadas às variáveis ambientais e, menos fortemente, em relação ao tamanho. Aves com status migratório apresentaram correlação significativa com pelo menos um dos eixos, mostrando-se mais fortemente associadas às manchas de FOM e áreas maiores. O uso de áreas abertas pelas aves teve uma associação mais marcante com os fragmentos e manchas de menor tamanho. Nectarívoros e frugívoros/folívoros tiveram uma associação mais forte com os fragmentos de FOD, enquanto as espécies onívoras foram mais bem representadas nas manchas de FOM. Aves de sub-bosque foram significativamente relacionadas aos fragmentos de FOD e áreas de menor tamanho e aves que habitam o dossel estiveram correlacionadas principalmente às manchas de FOM. A partir dos resultados obtidos nas áreas estudadas na Mata Atlântica do sul do Brasil, onde os maiores remanescentes florestais tem menos de 200 ha, observamos que mesmo as áreas de menor tamanho podem ser importantes para a manutenção das comunidades de aves. Conhecer as respostas da fauna às modificações ambientais impostas pela intervenção humana torna-se fundamental para o planejamento de estratégias de manejo e conservação. / Many issues in ecology, conservation biology and biogeography are related to the effects of natural or anthropogenic processes over fragments, patches and islands. Nowadays, the evaluation of fragments in a landscape context, as elements of a matrix including not only fragments of different size and continuous forest, but also other elements of landscape structure, is becoming increasingly important in order to understand fragmentation effects and to propose effective conservation actions. Furthermore, when comparing formations with distinct environmental attributes, even when geographically near, it is necessary to consider organization levels related to species and populations attributes to understand the mechanisms that drive some species to extinction and other to success. Some attributes may be used as ecological indicators that reflect environmental pressures on species biology, at the same time that reflect relationships with organism functions. In the first chapter of this thesis, we evaluated how methodological and analytical approaches may influence the results of bird studies and the relationship between species richness/abundance and the size of forest fragments and patches. We observed that the relationship between species richness and size of areas was affected by the standardization of the time counting. Species number increased significantly in patches of Mixed Ombrophilous Forest when counting was performed with unlimited radius. When considering bird records computed in each sampling minute, we found that with nine minutes of sampling the most part of the species and contact records had already been obtained. In the second chapter we investigated if rarefied richness and abundance of birds reduce with decreasing area of fragments at Atlantic Ombrophilous Dense Forest. At the same time, we aimed to understand how bird assembly is influenced by attributes of vegetation within fragments and around them. Our results indicated that species abundance in each fragment was significant related to the fragment size, with the highest abundance in the largest fragments. The analyses carried out between mean rarefied richness and the selected environmental variables showed that patch variables were more important than variables related to vegetation structure within fragments. In the third chapter we evaluated how bird communities are influenced by fragmentation in areas originally forested and by nucleation processes in areas of mosaic of patchy forests, in Atlantic Ombrophilous Dense Forest (AODF) and Mixed Ombrophilous Forest (MOF), respectively. MOF and AODF areas were aggregated when related to environmental variables and, less strongly, in relation to size. Migrant birds had significant correlation with at least one ordination axis, showing a stronger association with MOF patches and largest areas. The use of open areas was markedly associated with smaller fragments and patches. Nectarivores and folivores/frugivores had a strong association with AODF fragments, while omnivore species were better represented in MOF patches. Understory birds were significantly related to AODF fragments and small areas, while canopy birds were correlated mainly to MOF patches. Considering our results obtained in Atlantic Forest in southern Brazil – where the largest forest remnants have less than 200 ha, we observed that even small areas may be important for the conservation of bird communities. The knowledge about wildlife responds to environmental modifications made by humans is critical for management and conservation planning in this scenario.

Made available in DSpace on 2016-12-12T20:12:32Z (GMT). No. of bitstreams: 1 PGEF15MA042.pdf: 1375553 bytes, checksum: f93a1c6bfaca22be5ac8fae40f160f3c (MD5) Previous issue date: 2015-06-16 / The present dissertation aimed to understand the functional, phylogenetic and the floristic-structural organization of the tree component, along a topographic gradient in a forest in Upper Uruguay River region. Information of this nature is critical for the definition of conservation plans, since it allows inferences about the ecological strategies of species to occupy different environmental conditions. For this, 50 10 × 20 m permanent plots were allocated along different topographic sectors (lower, middle, upper), where all individual tree species with circumference at breast height (cbh) equal to or greater than 15.7 cm had the cbh measured, the height estimated and the botanical identity determined. For the 20 most abundant species, the following functional traits were determined: basic wood density, maximum potential height, leaf area, specific leaf area,leaves lifespan, regeneration and dispersal guilds. The floristic-structural organization was analyzed through Shannon (H ) and Pielou (J) Indices, NMDS (Non-metric Multidimensional Scaling), Permutational Multivariate Analysis of Variance (PERMANOVA), Indicator Species Analyzes and contingency table. The functional traits were analyzed by Rao Q Index and community weight means values for plots (CWM). The phylogenetic structure was 18 determined by Mean Pairwise Distance analysis and K statistic of phylogenetic signal. The functional and phylogenetic structures were compared among sectors by Analysis of Variance, for normal distributions, or Kruskal-Wallis, for non-normal distributions, followed by post hoc tests. The results demonstrated that the tree component showed floristic-structural and ecological strategies variations in function of topographic sectors. The taxonomic, functional and phylogenetic diversity was lower in the lower sector, with predominance of trees with low wood density, large leaves, with smaller values of specific leaf area, than sectors situated at upper positions in the terrain. It was concluded that the topography represented an important source of heterogeneity of the tree component in small spatial scale. The capacity of species occupy different sectors in relief was influenced by their functional traits, reflecting different ecological strategies, which are partly phylogenetically conserved / A presente dissertação teve como principal objetivo entender a organização florística-estrutural, funcional e filogenética do componente arbóreo, ao longo de um gradiente topográfico, em uma floresta na região do Alto Rio Uruguai. Informações desta natureza são fundamentais para a definição de planos de conservação, uma vez que permitem inferências sobre as estratégias ecológicas das espécies para se desenvolverem em condições ambientais distintas.Para isto, foram alocadas 50 parcelas permanentes de 10 × 20m, ao longo de diferentes setores topográficos (inferior, intermediário e superior), onde todos os indivíduos arbóreos com circunferência na altura do peito (CAP) maior ou igual a 15,7 cm, tiveram a CAP medida, a altura estimada e identidade botânica determinada. Para as 20 espécies mais abundantes, foram determinados os seguintes atributos funcionais: densidade básica da madeira, altura máxima potencial, área foliar, área foliar específica, regime de renovação foliar, guildas de regeneração e de dispersão.A organização florística-estrutural foi analisada por meio dos índices de Shannon (H ) e de Pielou (J), NMDS (Escalonamento Multidimensional Não-métrico), Análise de Variância Multivariada Permutacional (PERMANOVA), Análise de Espécies Indicadoras e tabela de contingência. Os atributos funcionais foram analisados por meio do índice de Rao Q e os valores médios dos atributos ponderados para comunidade em cada parcela (CWM). A estrutura filogenética foi determinada por meio da análise de Mean Pairwise Distance (MPD) e pela estatística K para sinal filogenético de atributos funcionais. As estruturas funcional e filogenética foram comparadas entre setores por meio de Análise de Variância, para distribuição normal, ou Kruskal-Wallis, para distribuições não-normais, com testes post hoc.Os resultados demonstraram que o componente arbóreo apresentou variações florísticas-estruturais e de estratégias ecológicas em função dos setores topográficos. A diversidade taxonômica, funcional e filogenética foi menor no Setor inferior, que apresentou a predominância indivíduos com baixa densidade da madeira, folhas maiores e com menor área foliar específica, em relação as posições mais elevadas no relevo. Conclui-se que a topografia representou uma importante fonte de heterogeneidade do componente arbóreo em pequena escala espacial. A capacidade das espécies ocuparem diferentes setores no relevo foi influenciada por seus atributos funcionais, refletindo diferentes estratégias ecológicas, que, em parte são filogeneticamente conservadas

Muitas questões ligadas à ecologia, biologia da conservação e biogeografia estão relacionadas aos efeitos de processos naturais ou antropogênicos sobre fragmentos, manchas ou ilhas. Na atualidade, para se compreender os efeitos da fragmentação e se propor medidas efetivas de conservação, torna-se cada vez mais importante analisar os fragmentos como parte de uma matriz que inclui não somente diferentes tamanhos de remanescentes e de floresta contínua, mas também outros elementos da estrutura da paisagem. Para tanto, quando comparamos formações com diferentes características ambientais, mesmo que inseridas em regiões próximas, para entendermos os mecanismos que levam ao desaparecimento de algumas espécies e beneficiam outras, é necessário considerar níveis de organização relacionados a atributos de espécies e populações. Determinados atributos podem ser indicadores ecológicos relevantes de pressões exercidas pelo ambiente na biologia das espécies, ao mesmo tempo em que demonstram relação com a função do organismo. Essa tese, em seu primeiro capítulo avalia como questões metodológicas e analíticas podem influenciar os resultados de estudos avifaunísticos e a relação da riqueza e abundância de espécies com o tamanho de fragmentos e manchas florestais. Observamos que a relação entre a riqueza e o tamanho das áreas foi afetada pela padronização do tempo de contagem por área. A utilização de contagens com raio ilimitado fez com que o número de espécies aumentasse significativamente somente nas maiores manchas de Floresta Ombrófila Mista. Considerando-se o acréscimo de registros obtidos em cada minuto de contagem, constatou-se que com nove minutos já haviam sido obtidos grande parte dos registros de espécies e contatos de aves. No segundo capítulo, foi investigado se a riqueza rarefeita e a abundância de aves diminuem com o decréscimo da área de fragmentos de Floresta Ombrófila Densa. Paralelamente buscou-se compreender como a avifauna é influenciada por características da vegetação dos fragmentos e do entorno destes. A abundância obtida em cada fragmento esteve significativamente relacionada ao tamanho das áreas, sendo mais elevada nos fragmentos maiores. As análises relacionando a riqueza rarefeita média dos fragmentos e as variáveis ambientais selecionadas mostraram que as variáveis de mancha foram mais importantes que as variáveis relacionadas à estrutura da vegetação no interior dos fragmentos. No terceiro capítulo, avaliamos como as comunidades de aves respondem diferentemente à fragmentação em regiões originalmente florestadas e à existência de mosaicos em regiões com manchas florestais (capões), situações características das regiões onde estão inseridas, respectivamente, as Florestas Ombrófilas Densa (FOD) e Mista (FOM). As áreas de FOM e FOD foram agrupadas quando relacionadas às variáveis ambientais e, menos fortemente, em relação ao tamanho. Aves com status migratório apresentaram correlação significativa com pelo menos um dos eixos, mostrando-se mais fortemente associadas às manchas de FOM e áreas maiores. O uso de áreas abertas pelas aves teve uma associação mais marcante com os fragmentos e manchas de menor tamanho. Nectarívoros e frugívoros/folívoros tiveram uma associação mais forte com os fragmentos de FOD, enquanto as espécies onívoras foram mais bem representadas nas manchas de FOM. Aves de sub-bosque foram significativamente relacionadas aos fragmentos de FOD e áreas de menor tamanho e aves que habitam o dossel estiveram correlacionadas principalmente às manchas de FOM. A partir dos resultados obtidos nas áreas estudadas na Mata Atlântica do sul do Brasil, onde os maiores remanescentes florestais tem menos de 200 ha, observamos que mesmo as áreas de menor tamanho podem ser importantes para a manutenção das comunidades de aves. Conhecer as respostas da fauna às modificações ambientais impostas pela intervenção humana torna-se fundamental para o planejamento de estratégias de manejo e conservação. / Many issues in ecology, conservation biology and biogeography are related to the effects of natural or anthropogenic processes over fragments, patches and islands. Nowadays, the evaluation of fragments in a landscape context, as elements of a matrix including not only fragments of different size and continuous forest, but also other elements of landscape structure, is becoming increasingly important in order to understand fragmentation effects and to propose effective conservation actions. Furthermore, when comparing formations with distinct environmental attributes, even when geographically near, it is necessary to consider organization levels related to species and populations attributes to understand the mechanisms that drive some species to extinction and other to success. Some attributes may be used as ecological indicators that reflect environmental pressures on species biology, at the same time that reflect relationships with organism functions. In the first chapter of this thesis, we evaluated how methodological and analytical approaches may influence the results of bird studies and the relationship between species richness/abundance and the size of forest fragments and patches. We observed that the relationship between species richness and size of areas was affected by the standardization of the time counting. Species number increased significantly in patches of Mixed Ombrophilous Forest when counting was performed with unlimited radius. When considering bird records computed in each sampling minute, we found that with nine minutes of sampling the most part of the species and contact records had already been obtained. In the second chapter we investigated if rarefied richness and abundance of birds reduce with decreasing area of fragments at Atlantic Ombrophilous Dense Forest. At the same time, we aimed to understand how bird assembly is influenced by attributes of vegetation within fragments and around them. Our results indicated that species abundance in each fragment was significant related to the fragment size, with the highest abundance in the largest fragments. The analyses carried out between mean rarefied richness and the selected environmental variables showed that patch variables were more important than variables related to vegetation structure within fragments. In the third chapter we evaluated how bird communities are influenced by fragmentation in areas originally forested and by nucleation processes in areas of mosaic of patchy forests, in Atlantic Ombrophilous Dense Forest (AODF) and Mixed Ombrophilous Forest (MOF), respectively. MOF and AODF areas were aggregated when related to environmental variables and, less strongly, in relation to size. Migrant birds had significant correlation with at least one ordination axis, showing a stronger association with MOF patches and largest areas. The use of open areas was markedly associated with smaller fragments and patches. Nectarivores and folivores/frugivores had a strong association with AODF fragments, while omnivore species were better represented in MOF patches. Understory birds were significantly related to AODF fragments and small areas, while canopy birds were correlated mainly to MOF patches. Considering our results obtained in Atlantic Forest in southern Brazil – where the largest forest remnants have less than 200 ha, we observed that even small areas may be important for the conservation of bird communities. The knowledge about wildlife responds to environmental modifications made by humans is critical for management and conservation planning in this scenario.

La disponibilité en eau est le principal facteur limitant le fonctionnement des écosystèmes méditerranéens. Des sècheresses plus marquées ou plus fréquentes pourraient avoir d'importantes répercussions sur l'activité et la diversité de la faune du sol qui régule la décomposition des litières et le cycle des nutriments. Dans cette thèse j'ai étudié expérimentalement les interactions entre une modification des apports en eau et l'impact de la macrofaune détritivore sur les processus de décomposition en conditions méditerranéennes. Dans une première partie, j'ai étudié l'effet d'une espèce de détritivore très abondante localement, Ommatoiuus sabulosus, sur la décomposition des litières d'arbustes de garrigue. Une expérience d'un mois en microcosmes a permis d'étudier ses effets directs (via la consommation de litière) et indirects (via l'activité microbienne dans ses fèces) sur la perte de masse des litières et les communautés microbiennes à deux niveaux d'humidité contrastés. Dans une autre expérience d'un an sur le terrain, la mise en place de sachets de litières et de fèces à deux profondeurs dans un sol de garrigue a permis d'étudier les effets d'Ommatoiulus à plus long terme. Les principaux résultats montrent que sa consommation de litière est moins affectée par la sècheresse que la décomposition microbienne, mais que, à court terme, Ommatoiulus ne stimule pas la minéralisation de la matière organique, quelles que soient les conditions d'humidité. En revanche, à plus long terme, Ommatoiulus peut accélérer la décomposition de certaines litières comme le chêne kermès, puisque des fèces issues de cette litière déposées à la surface du sol pendant un an perdent plus de masse que de la litière non-consommée. Cette stimulation semble liée à un lessivage plus important des composés organiques solubles dans les fèces et ne se produit qu'à la surface du sol. En profondeur, où l'humidité du sol est plus favorable à la décomposition, la perte de masse des fèces augmente. Ce résultat suggère qu'en facilitant l'enfouissement de la matière organique dans le sol, les détritivores peuvent accélérer la décomposition.Dans une seconde partie, j'ai cherché à évaluer l'importance de la diversité fonctionnelle des litières et des détritivores pour le processus de décomposition. Grâce à une approche basée sur les traits, des assemblages d'espèces représentant un fort gradient de dissimilarité fonctionnelle mais ayant une richesse spécifique constante, ont été créés pour étudier la réponse de la relation diversité-fonction à la sécheresse. Les résultats de cette expérience menée à l'Ecotron de Montpellier, montrent que la dissimilarité fonctionnelle des litières et des détritivores explique jusqu'à 20% de la variation observée dans plusieurs processus clefs du fonctionnement du sol, tels que la perte de masse des litières et le lessivage du carbone et de l'azote dans le sol superficiel. Toutefois, les effets de l'identité des espèces présentes aux deux niveaux trophiques restent plus importants que ceux de la dissimilarité fonctionnelle. Bien que la sécheresse influence fortement les processus étudiés, les relations diversité-fonction ne sont pas modifiées par un changement de la disponibilité en eau. Cependant, les assemblages d'espèces les plus performants en conditions d'humidité favorables sont aussi les plus fortement affectés par la sécheresse, ce qui suggère qu'il existe un compromis entre l'efficacité des organismes du sol et leur capacité à résister à une perturbation. / Water availability is a major limiting factor for the functioning of Mediterranean ecosystems. More pronounced drought could severely impact soil fauna activity and diversity that could in turn affect litter decomposition and nutrient cycling. In my PhD thesis I investigated experimentally the interactions between changing water availability and detritivorous macrofauna on decomposition and associated processes in a “garrigue”, a typical Mediterranean woody shrub dominated ecosystem.In the first part of my thesis, I studied the impact of Ommatoiulus sabulosus, an abundant diplopod species in garrigue ecosystems, on shrub litter decomposition. During a one month experiment, I studied the direct (litter consumption) and indirect (microbial activity in feces) effects of this detritivore on litter mass loss and microbial communities under two contrasted moisture levels. In a different experiment, I placed litterbags filled with litter or feces in the field at the soil surface or at 5cm soil depth during one year in order to study the long term impact of Ommatoiulus on decomposition. A key result was that detritivores maintain litter consumption in dry conditions when microbial driven decomposition drastically dropped. However, this detritivore effect do not lead to an overall increased organic matter mineralization irrespective of moisture conditions, at least in the short term. In contrast, under field conditions and over a longer time period, Ommatoiulus increases decomposition of certain species such as Quercus coccifera, since feces from this species decomposes faster than un-ingested litter after one year at the soil surface. This stimulation is likely due to a higher leaching of soluble compound in feces. Moreover, in depth feces decomposition increases relative to that of intact leaf litter, possibly indicating that more favorable soil humidity is more favorable to decomposition. Collectively, my results suggest that detritivores can strongly increase decomposition by transforming leaf litter into feces of different organic matter quality, and by facilitating the transfer of organic matter into the soil.In the second part, I evaluated the importance of functional dissimilarity of leaf litter and detritivores on decomposition processes. Using a trait based approach, species assemblages were constructed in order to obtain a gradient of functional dissimilarity of both, leaf litter and detritivore communities, while keeping species numbers constant. The different communities were kept under controlled conditions at the European Ecotron in Montpellier to study the effect of changing functional dissimilarity on process rates at two different moisture conditions. I found that detritivore and litter functional dissimilarity explain up to 20 % of the observed variation for several key soil processes including litter mass loss and the leaching of dissolved organic carbon and nitrogen from top soil. However, effects of species identity at both trophic levels have a larger impact on process rates than functional dissimilarity. In general, drought strongly affects soil processes but does not alter the diversity-function relationship. Species assemblages resulting in highest process rates at favorable moisture level are also the most negatively affected by drought, suggesting a tradeoff between the efficiency of soil organisms and their ability to resist perturbation.

Muitas questões ligadas à ecologia, biologia da conservação e biogeografia estão relacionadas aos efeitos de processos naturais ou antropogênicos sobre fragmentos, manchas ou ilhas. Na atualidade, para se compreender os efeitos da fragmentação e se propor medidas efetivas de conservação, torna-se cada vez mais importante analisar os fragmentos como parte de uma matriz que inclui não somente diferentes tamanhos de remanescentes e de floresta contínua, mas também outros elementos da estrutura da paisagem. Para tanto, quando comparamos formações com diferentes características ambientais, mesmo que inseridas em regiões próximas, para entendermos os mecanismos que levam ao desaparecimento de algumas espécies e beneficiam outras, é necessário considerar níveis de organização relacionados a atributos de espécies e populações. Determinados atributos podem ser indicadores ecológicos relevantes de pressões exercidas pelo ambiente na biologia das espécies, ao mesmo tempo em que demonstram relação com a função do organismo. Essa tese, em seu primeiro capítulo avalia como questões metodológicas e analíticas podem influenciar os resultados de estudos avifaunísticos e a relação da riqueza e abundância de espécies com o tamanho de fragmentos e manchas florestais. Observamos que a relação entre a riqueza e o tamanho das áreas foi afetada pela padronização do tempo de contagem por área. A utilização de contagens com raio ilimitado fez com que o número de espécies aumentasse significativamente somente nas maiores manchas de Floresta Ombrófila Mista. Considerando-se o acréscimo de registros obtidos em cada minuto de contagem, constatou-se que com nove minutos já haviam sido obtidos grande parte dos registros de espécies e contatos de aves. No segundo capítulo, foi investigado se a riqueza rarefeita e a abundância de aves diminuem com o decréscimo da área de fragmentos de Floresta Ombrófila Densa. Paralelamente buscou-se compreender como a avifauna é influenciada por características da vegetação dos fragmentos e do entorno destes. A abundância obtida em cada fragmento esteve significativamente relacionada ao tamanho das áreas, sendo mais elevada nos fragmentos maiores. As análises relacionando a riqueza rarefeita média dos fragmentos e as variáveis ambientais selecionadas mostraram que as variáveis de mancha foram mais importantes que as variáveis relacionadas à estrutura da vegetação no interior dos fragmentos. No terceiro capítulo, avaliamos como as comunidades de aves respondem diferentemente à fragmentação em regiões originalmente florestadas e à existência de mosaicos em regiões com manchas florestais (capões), situações características das regiões onde estão inseridas, respectivamente, as Florestas Ombrófilas Densa (FOD) e Mista (FOM). As áreas de FOM e FOD foram agrupadas quando relacionadas às variáveis ambientais e, menos fortemente, em relação ao tamanho. Aves com status migratório apresentaram correlação significativa com pelo menos um dos eixos, mostrando-se mais fortemente associadas às manchas de FOM e áreas maiores. O uso de áreas abertas pelas aves teve uma associação mais marcante com os fragmentos e manchas de menor tamanho. Nectarívoros e frugívoros/folívoros tiveram uma associação mais forte com os fragmentos de FOD, enquanto as espécies onívoras foram mais bem representadas nas manchas de FOM. Aves de sub-bosque foram significativamente relacionadas aos fragmentos de FOD e áreas de menor tamanho e aves que habitam o dossel estiveram correlacionadas principalmente às manchas de FOM. A partir dos resultados obtidos nas áreas estudadas na Mata Atlântica do sul do Brasil, onde os maiores remanescentes florestais tem menos de 200 ha, observamos que mesmo as áreas de menor tamanho podem ser importantes para a manutenção das comunidades de aves. Conhecer as respostas da fauna às modificações ambientais impostas pela intervenção humana torna-se fundamental para o planejamento de estratégias de manejo e conservação. / Many issues in ecology, conservation biology and biogeography are related to the effects of natural or anthropogenic processes over fragments, patches and islands. Nowadays, the evaluation of fragments in a landscape context, as elements of a matrix including not only fragments of different size and continuous forest, but also other elements of landscape structure, is becoming increasingly important in order to understand fragmentation effects and to propose effective conservation actions. Furthermore, when comparing formations with distinct environmental attributes, even when geographically near, it is necessary to consider organization levels related to species and populations attributes to understand the mechanisms that drive some species to extinction and other to success. Some attributes may be used as ecological indicators that reflect environmental pressures on species biology, at the same time that reflect relationships with organism functions. In the first chapter of this thesis, we evaluated how methodological and analytical approaches may influence the results of bird studies and the relationship between species richness/abundance and the size of forest fragments and patches. We observed that the relationship between species richness and size of areas was affected by the standardization of the time counting. Species number increased significantly in patches of Mixed Ombrophilous Forest when counting was performed with unlimited radius. When considering bird records computed in each sampling minute, we found that with nine minutes of sampling the most part of the species and contact records had already been obtained. In the second chapter we investigated if rarefied richness and abundance of birds reduce with decreasing area of fragments at Atlantic Ombrophilous Dense Forest. At the same time, we aimed to understand how bird assembly is influenced by attributes of vegetation within fragments and around them. Our results indicated that species abundance in each fragment was significant related to the fragment size, with the highest abundance in the largest fragments. The analyses carried out between mean rarefied richness and the selected environmental variables showed that patch variables were more important than variables related to vegetation structure within fragments. In the third chapter we evaluated how bird communities are influenced by fragmentation in areas originally forested and by nucleation processes in areas of mosaic of patchy forests, in Atlantic Ombrophilous Dense Forest (AODF) and Mixed Ombrophilous Forest (MOF), respectively. MOF and AODF areas were aggregated when related to environmental variables and, less strongly, in relation to size. Migrant birds had significant correlation with at least one ordination axis, showing a stronger association with MOF patches and largest areas. The use of open areas was markedly associated with smaller fragments and patches. Nectarivores and folivores/frugivores had a strong association with AODF fragments, while omnivore species were better represented in MOF patches. Understory birds were significantly related to AODF fragments and small areas, while canopy birds were correlated mainly to MOF patches. Considering our results obtained in Atlantic Forest in southern Brazil – where the largest forest remnants have less than 200 ha, we observed that even small areas may be important for the conservation of bird communities. The knowledge about wildlife responds to environmental modifications made by humans is critical for management and conservation planning in this scenario.