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  • About
  • The Global ETD Search service is a free service for researchers to find electronic theses and dissertations. This service is provided by the Networked Digital Library of Theses and Dissertations.
    Our metadata is collected from universities around the world. If you manage a university/consortium/country archive and want to be added, details can be found on the NDLTD website.
1

Bacterioplankton population dynamics in a changing ocean

Lindh, Markus V. January 2014 (has links)
Bacterioplankton is characterized by high diversity, short generation times and rapid turnover. Despite their small size, these numerous microorganisms are a fundamental piece of aquatic ecosystems by channeling carbon to higher trophic levels through dissolved organic matter utilization. Yet, several gaps remain in our knowledge and understanding of bacterioplankton populations regarding detailed temporal dynamics, and mechanisms determining biogeographical patterns and potential responses to climate change. The aim of this thesis was to examine responses in bacterioplankton community composition and function when challenged by natural and anthropogenically-induced change in environmental conditions. High temporal resolution analysis of bacterioplankton population dynamics in the Baltic Sea indicated detailed seasonal responses. It also showed a similar but wide spectrum of niche differentiation patterns within several major bacterial groups. Analysis of geographic distributions of marine bacterial populations revealed bimodal occupancy-frequency patterns in bacterial communities, indicating that the presence of many locally rare taxa along with a few locally abundant taxa were explained by stochastic variation in colonization and extinction rates. Experimental manipulations with natural marine bacterioplankton assemblages revealed both specialist and generalist strategies in utilizing specific dissolved organic carbon compounds. When subjected to experimentally increased sea surface temperatures, lowered pH and additions of terrigenous carbon, some populations decreased in relative abundance while others were stable; concomitantly, many populations increased in relative abundance. Shifts in bacterial community composition were shown to correlate with changes in community functioning, but detection of such correlations depended largely on the detail of phylogenetic analysis and successional stage of the communities. The results in this thesis suggest that both natural and anthropogenically-induced changes in environmental conditions promote simultaneous adjustment and replacement of bacterial populations tightly linked with metabolic plasticity. These trade-offs play a significant role for understanding the relationship between bacterioplankton population dynamics and potential shifts in carbon cycling properties. We also show the importance of regional effects in shaping bacterial community composition, crucial for interpreting bacterioplankton distribution patterns. In conclusion, this thesis emphasizes the critical importance of connecting analysis of bacterioplankton population dynamics with examination of ecological mechanisms to improve our understanding of factors that regulate the distribution and activity of distinct bacterioplankton populations. / Hälften av all fotosyntes på vår planet utförs av växtplankton. De producerar organiskt material som utgör grunden för näringskedjan i havet. Ungefär hälften av det organiska material som produceras av växtplankton utnyttjas inte direkt, utan omsätts istället av bakterieplankton som lever och växer fritt i vattenmassan eller på olika partiklar. Bakterieplankton spelar därmed en nyckelroll i ekosystemet genom sin konsumtion av organiskt kol som för energi högre upp i näringskedjan. Trots deras nyckelroll i akvatiska miljöer vet vi fortfarande mycket lite om bakteriernas detaljerade säsongsmönster, mekanismer bakom rumsliga mönster och hur olika populationer kan komma att svara på klimatförändringar. Målet med denna avhandling var att undersöka hur specifika populationers dynamik och ekosystemfunktion påverkas av naturliga eller klimatorsakade förändringar i havsmiljön. Våra resultat av högupplöst säsongsbunden dynamik i Östersjöns bakteriesamhälle avslöjar en liknande bred uppdelning av ekologiska strategier inom varje större grupp av bakterier, både i relativ abundans och temporal fördelning. Utbredning i rum och tid av många lokalt ovanliga populationer jämfört med få lokalt vanliga populationer förklarades genom stokastisk variation i kolonisations- och utdöendehastigheter. Vidare tyder experimentella studier med tillsatser av olika kolkällor på att marina bakterier har olika ekologiska strategier, där populationer är specialister eller generalister i utnyttjandet av enskilda kolkällor. Med hjälp av experiment med naturliga bakteriesamhällen bekräftade vi tydliga temperatureffekter på bakteriesamhällets sammansättning, och en mindre effekt av lägre pH - som dock tillsammans med förhöjd temperatur bidrog till en tydlig synergistisk effekt på artsammansättningen. Ökad temperatur tillsammans med tillsats av terrestert kol gav också en stor effekt på bakteriesamhällets struktur och ekosystemfunktion och pekar på en potentiellt viktig påverkan av ökad framtida nederbörd och avrinning från vattendrag till havet. Samtliga tre experiment med fokus på klimatpåverkan bekräftade förekomsten av populationer som försvann eller minskade i relativ abundans vid klimatpåverkan (känslighet), medan andra var stabila (resistens). Samtidigt svarade många populationer positivt på klimatorsakade förändringar i havsmiljön och ökade i relativ abundans (respons) samtidigt som bakteriernas ekosystemfunktion påverkades positivt. Sammanfattningsvis visar denna avhandling att vissa nya bakteriepopulationer kan etablera sig och ersätta andra samtidigt som vissa befintliga populationer anpassar sin livsstrategi och ekologi till förändringar i havsmiljön. Vi visar också vikten av regionala effekter, d.v.s. kolonisation och utdöende, för bakteriesamhällets struktur, viktigt för tolkningen av biogeografiska mönster och den genomiska potentialen hos specifika populationer. Denna avhandling poängterar därmed betydelsen av att koppla studier av ekologiska mekanismer till både rumsliga och temporala spridningsmönster hos bakterier och till populationers kapacitet att svara på och anpassa sig till förändringar i havsmiljön.
2

A inclusão da abundância na diversidade funcional aumenta o seu poder de previsão?: teste em uma comunidade de cerrado / Does the inclusion of abundance in functional diversity increase its predictive power?: test in a brazilian savanna

Miatto, Raquel Carolina 04 March 2011 (has links)
Made available in DSpace on 2016-06-02T19:31:55Z (GMT). No. of bitstreams: 1 3610.pdf: 381494 bytes, checksum: 801cdcfd0eb82f33c299301532707d13 (MD5) Previous issue date: 2011-03-04 / Financiadora de Estudos e Projetos / Assuming that traditional measures of diversity are rough predictors of functioning and that measures that incoporate information about functional traits are better predictors than the traditional ones, functional diversity measures are intensely used nowadays. However, whether the inclusion of functional traits in biodiversity estimation indeed increases our predictive power is rarely tested. Moreover, dendrogram-based functional diversity measures usually do not take abundance into account. By sampling a southern cerrado site in Brazil, we tested whether, when compared to traditional diversity measures, the inclusion of functional traits leads to an increased predictive power in relating functional diversity to community functioning. We demonstrated that it is possible to include abundance in dendrogram-based functional diversity, and tested whether, when compared to functional diversity as usually calculated, the inclusion of abundance leads to an increased predictive power. No diversity measure was related to community functioning, including functional diversity. Nevertheless, we showed that it is possible to include abundance in dendrogram16 based functional diversity measures, even if most indices were correlated with each other. / Assumindo que medidas de diversidade tradicionais são estimativas grosseiras do funcionamento de comunidades e que medidas que incorporam informações dos traços funcionais são melhores do que as tradicionais, medidas de diversidade funcional são bastante utilizadas atualmente. Contudo, raramente é testado se, de fato, a inclusão dos traços funcionais em estimativas de biodiversidade aumenta nosso poder preditivo. Além disso, medidas de diversidade funcional baseadas em dendrogramas geralmente não consideram a abundância. Em uma área de cerrado no sudeste brasileiro, testamos se a inclusão dos traços funcionais leva a um aumento no poder preditivo ao relacionar a diversidade funcional ao funcionamento da comunidade quando comparada a medidas de diversidade tradicionais. Demonstramos que é possível incluir a abundância em medidas de diversidade funcional baseadas em dendrogramas e testamos se a inclusão da abundância eleva o poder preditivo da diversidade funcional em comparação à comumente calculada. Nenhuma medida de diversidade relacionou-se ao funcionamento da comunidade, inclusive a diversidade funcional. Porém, mostramos que é possível incluir a abundância em medidas de diversidade funcional baseadas em dendrogramas, apesar da maioria dos índices estar correlacionada.
3

Processes involved in the functioning of large mammal communities : the role of the African elephant in the ecology of predator-prey relationships / Processus impliqués dans le fonctionnement des communautés de grands mamifères : le rôle de l'éléphant Africain dans l'écologie des relations prédateurs-proie

Ferry, Nicolas 06 April 2018 (has links)
Les communautés écologiques sont connues pour être des systèmes complexes composés de multiple espèces entrant en interaction les unes avec les autres. De nombreux modèles théoriques ont été développés pour étudier les communautés. Certains ont souligné l'importance des effets indirects que les espèces pouvaient avoir les unes sur les autres, tels que les chaînes d'interactions et les modifications d'interactions (par modification du trait d'une des espèces en interaction ou de l'environnement où se déroule l'interaction). Bien que la science expérimentale vienne confirmer le rôle fondamental que pourrait avoir ces effets indirects, peu d'études à l'échelle des communautés en milieu naturel ont été conduites, et encore moins chez les grands mammifères. Le Parc National de Hwange, au Zimbabwe, est un écosystème de savane arborée semiaride caractérisé par une quasi-absence d'eau de surface naturelle (point d'eau, rivière) pendant la saison sèche, et ce n'est qu'avec la création de points d'eau artificiels pompés que la richesse spécifique des communautés de grands mammifères et les fortes abondances animales sont maintenues. De plus, cet écosystème a la remarquable particularité à la fois d'abriter l'une des plus fortes densités d'éléphants, et d'être considéré comme l'un des bastions africains pour sa population de lions. Le lion est connu comme étant un chasseur à l'affût, utilisant les éléments de son habitat (fourrés, souches, hautes herbes, etc.) pour se rapprocher au maximum de sa proie et lui bondir dessus par surprise. De plus, cette espèce semble profiter de l'agrégation des herbivores aux points d'eau pendant la saison sèche pour chasser autour de ces points d'eau. L'éléphant d'Afrique quant à lui est capable d'aménager son milieu et est ainsi susceptible de favoriser l'accès à certaines ressources pour les autres espèces, telles que des abris ou au contraire une meilleure visibilité. En revanche, de par sa masse corporelle exceptionnelle et son régime alimentaire généraliste, il est possible qu'il soit un compétiteur clé pour les autres herbivores. Enfin, étant très nombreux dans l'écosystème étudié, nécessitant de grandes quantités d'eau, et devenant de plus en plus agressifs au fil de la saison sèche, les éléphants influencent l'utilisation des points d'eau par les autres herbivores. Cette thèse porte donc sur le rôle que peuvent avoir les éléphants sur les interactions trophiques entre les lions et leurs proies, via des mécanismes d'effets indirects. Différents axes de recherche sont abordés. Le premier porte sur l'effet des éléphants sur la communauté de grands herbivores aux points d'eau, et plus particulièrement comment ils peuvent influencer leur distribution spatiale, et à terme leur vulnérabilité vis-à-vis des prédateurs. Un évitement spatial des éléphants par les autres herbviores en début de saison sèche suggère fortement que les éléphants sont de potentiels compétiteurs. Cependant, à la fin de la saison sèche, le phénomène s'inverse et certaines espèces d'herbivores se rapprochent fortement des éléphants. Deux scénarios portant sur les mécanismes pouvant expliquer ce patron ont été explorés, sans succès : une nécessité croissante d'accéder à de l'eau de meilleure qualité au niveau des pompes des points d'eau, et une augmentation du risque de prédation qui pourrait rendre les éléphants « attractifs » aux yeux des herbivores, les éléphants adultes étant invulnérables à la prédation et capables de les faire fuir par des comportements de harcèlement. Le deuxième axe de recherche porte sur l'effet des éléphants sur la distribution spatiale des herbivores à l'échelles du paysage et de l'habitat, et sur les conséquences possibles que cela peut avoir sur l'écologie spatiale des prédateurs. L'absence de ségrégation entre éléphants et herbivores ne supporte pas l'hypothèse d'un effet de compétition par exploitation, et l'investigation quant à l'effet sur les prédateurs n'a pas été poussée plus avant... [etc] / Species can indirectly affect other species and their interactions. The trophic interaction between a predator and its prey can be modified by the presence of a third species either through chain interactions (e.g. successive predation link) either through interaction modification. However, these indirect received few attentions in theorical modelling of food web, and fewer studies tried to explore this phenomenon at the scale of natural complex communities of large mammals. The role of the elephants as modifier of lion’s trophic interaction is explored in the semi-arid woodland savannah ecosystem of Hwange National Park, Zimbabwe. African elephants, as key competitor (male body mass ~ 4000 kg with aggressive behaviour) shape the behaviour of herbivores at waterholes results do not allow to state on the elephant mediation of lion trophic interaction at waterholes. In addition, elephants seem to facilitate the availability of food resources for impalas, possibly by increasing regrowth of shoots by breaking twigs and stem, as these last select habitats used by elephants. However, not effect of facilitation or competition were observed for the other herbivores, which lead to think that elephants do not influence lion trophic interaction in that way. Finally, by altering the physical environment (i.e. engineer species) the elephants affect the visibility and ambush sites for lions in the woody vegetation and ultimately seem to influence the lion kill site selection. This study suggests that indirect effects may act at the community level even if their observation and quantification are difficult in natural communities. Moreover, it supports the observation that it is important to take into account these indirect effects in order to have a thorough understanding and have a better ability to predict the consequences that disruptions may have on the structure and functioning of communities
4

Complementaridade de atributos reduz a decomposição de folhas em assembleias experimentais

Plazas Jiménez, Oscar Daniel 06 March 2018 (has links)
Submitted by Luciana Ferreira (lucgeral@gmail.com) on 2018-04-09T12:04:41Z No. of bitstreams: 2 Dissertação - Oscar Daniel Plazas Jiménez - 2018.pdf: 5372786 bytes, checksum: ccb044675f67ad084ee0892aa9c09bae (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) / Approved for entry into archive by Luciana Ferreira (lucgeral@gmail.com) on 2018-04-09T12:42:46Z (GMT) No. of bitstreams: 2 Dissertação - Oscar Daniel Plazas Jiménez - 2018.pdf: 5372786 bytes, checksum: ccb044675f67ad084ee0892aa9c09bae (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) / Made available in DSpace on 2018-04-09T12:42:46Z (GMT). No. of bitstreams: 2 Dissertação - Oscar Daniel Plazas Jiménez - 2018.pdf: 5372786 bytes, checksum: ccb044675f67ad084ee0892aa9c09bae (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Previous issue date: 2018-03-06 / Theoretical and empirical evidence suggest that the mass ratio and the niche complementarity hypothesis are able to explain how species trait composition drives decomposition, an ecosystem process related with nutrient cycling and carbon sequestration. These two hypotheses are respectively related with the mean trait values of individuals present in a given community and its variance. Thus, it is impossible to test them adequately without experiments planned to approach these two aspects simultaneously. Although leaf functional traits influence decomposition process, an experimental design made to disentangle the unique, combined and strength of effects of the mechanisms proposed by MRH and NCH on decomposition should use leaf functional traits as predictors, but this has not yet been done. Here, we used two leaf nutritional traits (leaf nitrogen and magnesium-calcium content) and one structural trait (specific leaf area; SLA) from 20 species of seasonal dry forest to simulate leaf mixtures with different mean trait values (calculated as community-weighted mean; CWM) and variation of trait values (functional diversity; FD). For each trait, 40 leaf mixtures with different quasi-orthogonal combinations of CWM and FD were selected and left in the field to decompose for six months. We found that MRH mechanisms exert predominant role on decomposition where low and high CWM values were related with high biomass loss. Then, this add more evidence to support that the mean trait values of leaf mixtures are the main drivers of biomass loss. Contrary to previously evidences, trait complementarity does not increase efficiency and instead, an increasing in niche complementarity (high FD values) was followed by a decreasing in biomass loss efficiency. So, this question the importance of mechanism behind NCH on leaf decomposition. Additionally, we found that whereas nutritional traits were able to predict leaf decomposition, structural traits were not, in opposite to what is reported in the literature. Our results highlight that the mean resources quality of the leaf mixtures is the main driver of biomass loss and trait complementarity does not increase efficiency of biomass loss. Due to this, we proposed that in future studies mean resources quality of the leaf mixtures will be use as parameter to test and predict the possible impacts of human-induced changes on nutrient cycling and carbon sequestration. / Evidências empíricas e teóricas sugerem que os mecanismos propostos pelas hipóteses da razão da massa e da complementaridade de nicho (HRM e HCN respectivamente) são capazes de explicar como a composição de atributos funcionais das espécies determina a decomposição, um importante processo ecossistêmico relacionado com a ciclagem de nutrientes e o sequestro de carbono. Essas duas hipóteses estão relacionadas, respectivamente, com o valor médio dos atributos das espécies presentes em uma dada assembleia ou com a sua variação. Portanto, é impossível testar adequadamente essas duas hipóteses sem experimentos planejados para abordar simultaneamente esses dois aspectos, algo que é raramente feito. Os atributos funcionais foliares apresentam forte efeito nas taxas de decomposição em ecossistemas terrestres. Assim, um desenho experimental feito para separar a magnitude relativa dos efeitos únicos e combinados da HRM e HCN sobre a decomposição de folhas deve usar atributos foliares como preditores, mas até agora isso não foi feito. Neste trabalho, simulamos misturas de folhas com diferentes valores médios ponderados pela abundância (i.e., community weighted mean, CWM) de atributos e dispersão (i.e., functional diversity; FD) destes valores para dois atributos foliares nutricionais (conteúdo de nitrogênio e conteúdo de magnésio e cálcio) e um atributo estrutural (área foliar específica; SLA) quantificados para 20 espécies de floresta estacional semidecídua. Para cada atributo, selecionamos 40 misturas de folhas com diferentes combinações quase-ortogonais de CWM e FD que posteriormente foram deixadas durante seis meses no campo para decompor. No geral, encontramos que os mecanismos propostos pela HRM têm um papel predominante na decomposição, porque altos e baixos valores de CWM estiveram relacionados com uma alta perda de biomassa. Isso adiciona mais evidência à que os valores médios dos atributos das misturas de folhas são os principais determinantes da perda de biomassa. Porém, diferente do esperado, o aumento da complementaridade dos atributos das misturas de folhas (isto é, altos 8 valores de FD) levou à uma diminuição da eficiência da perda de biomassa e não à um aumento, como era esperado. Isso coloca em questão a importância dos mecanismos propostos pela HCN na decomposição de folhas. Adicionalmente, encontramos que os atributos nutricionais foliares foram capazes de predizer a perda de biomassa enquanto que, ao contrário do reportado na literatura, o atributo estrutural SLA não teve qualquer relação com a decomposição. Nossos resultados destacam que em misturas de folhas os valores médios do conteúdo de nutrientes estão associados com as taxas de decomposição e a complementaridade dos atributos não incrementou a eficiência na perda de biomassa. Portanto, nós propomos que em estudos futuros os valores médios do conteúdo de nutrientes das misturas de folhas sejam usados como parâmetros para testar e predizer os possíveis efeitos das mudanças na biodiversidade induzidos pelo homem sobre a ciclagem de nutrientes e o sequestro de carbono
5

What does a bioenergetic network approach tell us about the functioning of ecological communities?

Delmas, Eva 05 1900 (has links)
Les perturbations auxquelles font face les communautés écologiques, du fait des activités humaines, sont à l'origine de changements profonds dans ces communautés. Nombreuses caractéristiques des espèces sont altérées, de leur physiologie à leur occurrence même. Ces changements se répercutent sur la composition, la diversité et la structure des communautés, puisque les espèces n'interagissent pas tout le temps de la même manière en fonction des conditions. Prévoir le devenir de ces communautés émergentes, et des fonctions qu'elles soutiennent est un défi central de l'écologie et de nos sociétés. Différents cadres conceptuels ont été utilisés pour relever ce défi, basés sur différents mécanismes écologiques, et ont divergé en plusieurs domaines. D'un côté, l'analyse des chaînes trophiques utilise la consommation pour expliquer les effets de la diversité verticale (le nombre de niveaux trophiques) sur le fonctionnement, et de l'autre côté, les analyses biodiversité-fonctionnement lient compétition et effets de la diversité horizontale (la diversité au sein des niveaux trophiques isolés). Chacun de ces domaines a produit des résultats clés pour comprendre les conséquences fonctionnelles des changements de composition et diversité des communautés écologiques. Cependant, ils sont chacun basés sur différentes simplifications fortes des communautés. L'hypothèse qui sous-tend cette thèse est que la réconciliation en un même cadre de travail des résultats fondamentaux de ces champs conceptuels divergents, ainsi que des effets des changements de structure de la biodiversité, est une étape clé pour pouvoir améliorer notre compréhension du fonctionnement de communautés écologiques en changement. L'essor récent des méthodes d'analyse des réseaux trophiques, et des modèles permettant de simuler le fonctionnement de ces réseaux trophiques offre un cadre idéal pour cette réconciliation. En effet, les réseaux trophiques cartographient les échanges de matière entre toutes les espèces d'une communauté, permettant la mise en place d'interactions variées. Ils reflètent mieux la réalité complexe des communautés que les chaînes trophiques ou leurs niveaux trophiques isolés en intégrant notamment compétition et consommation. Un modèle ressource-consommateur bioénergétique classique, développé par Yodzis et Innes (1992), permet d'en simuler le fonctionnement, en intégrant des mécanismes et taux testés empiriquement. Au-delà d'utiliser ces outils, cette thèse se concentre aussi sur leur évaluation. Après un premier chapitre d'introduction, le second chapitre propose une plateforme ouverte, commune, solidement testée et efficace pour l'utilisation du modèle bioénergétique, permettant ainsi une synthèse plus rapide et aisée des résultats. Le troisième chapitre est une revue du corpus méthodologique d'analyse des réseaux trophiques, proposant une gamme de méthodes robustes et informatives, et soulignant leur domaine d'application et leurs limites. Enfin le quatrième chapitre met ce cadre méthodologique à l'épreuve. Dans ce chapitre, nous montrons l'existence d'une relation entre la complexité de la structure du réseau trophique des communautés et leur régime de fonctionnement, se traduisant par la réalisation de différentes prédictions issues de l'analyse des chaînes trophiques ou des analyses diversité-fonctionnement. Cette mise en évidence des conditions structurelles pour la réalisation de différentes prédictions nous permet de mieux comprendre quels mécanismes écologiques prédominent selon différentes conditions, dirigeant l'effet de la diversité sur le fonctionnement. / Human-driven disturbances are causing profound changes in ecological communities, as many characteristics of species are altered, from their physiology to their very occurrence. These changes affect the composition, diversity and structure of communities, since species do not always interact in the same way under different conditions. Predicting the fate of these emerging communities, and the functions they support, is a central challenge for ecology and our societies. Diverging conceptual frameworks have been used to address this challenge, based on different ecological mechanisms. On the one hand, food chain analysis uses consumption to explain the effects of vertical diversity (the number of trophic levels) on functioning, and on the other hand, biodiversity-functioning analyses link competition and the effects of horizontal diversity (diversity within isolated trophic levels). Each of these domains has produced key results for understanding the functional consequences of changes in the composition and diversity of ecological communities. However, they are each based on different strong simplifications of communities. The hypothesis underlying this thesis is that reconciling the fundamental results of these divergent conceptual fields, as well as the effects of changes in the structure of biodiversity, into a single framework is a key step towards improving our understanding of the functioning of changing ecological communities. The recent development of food web analysis and of models to simulate food webs functioning provides an ideal framework for this reconciliation. Food webs map the exchange of matter between all species in a community, allowing for a variety of interactions to take place. They better reflect the complex reality of communities than food chains or their isolated trophic levels, notably by integrating competition and consumption. A classical consumer-resource bioenergetic model developed by Yodzis and Innes (1992) specifically makes it possible to realistically simulate their functioning, using empirically tested mechanisms and rates. Beyond using these tools, this thesis focuses on their evaluation and implementation. After a first, introductory chapter, the second chapter proposes an open, common, well-tested and efficient platform for the use of the bioenergetic model, allowing a faster and easier synthesis of the results. The third chapter is a review of the methodological corpus for ecological networks analysis, outlining a range of robust and informative methods, and highlighting their scope and limitations. Finally, the fourth chapter puts this methodological framework to the test. In this chapter, we show the existence of a relationship between the complexity of communities' food-web structure and functioning regime, resulting in the realization of different predictions from food chain analysis or diversity-functioning analyses. This demonstration of the structural conditions for the realization of different predictions allows us to better understand which ecological mechanisms predominate under different conditions, directing the effect of diversity on functioning.

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