A field experiment from 2003 to 2005 at two sites examined the impacts of forage species and legume proportion on forage sward production. Grasses generally established rapidly and out-yielded swards high in legume content, although legumes did improve forage quality. Alfalfa was retained at greater relative biomass in mixed swards than swards containing clover. Legume persistence also varied depending on neighbouring grass species. A greenhouse study examined competitive interactions between Canada thistle (a common pasture weed), white clover and Kentucky bluegrass during establishment. Although thistle was most susceptible to intra-specific competition, and strongly affected forage yield, the latter also influenced weed biomass. Competitiveness of forages depended directly on soil medium, emphasizing the importance of abiotic factors on vegetation dynamics in mixed swards. / Rangeland and Wildlife Resources

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Unequal sex ratios are widespread in dioecious plants and understanding their cause is important to understanding fundamental aspects of their population dynamics, and yet what causes biased sex ratios in plants is still poorly understood. Competition experiments have been used in plants to predict the outcome of species interactions, but they have rarely been used to help explain sex ratio bias. This study used a response surface competition design to measure the relative competitive abilities of the sexes of the bryophyte Marchantia inflexa (a thallus liverwort of Marchantiaceae) to predict the outcome of competition before the onset of sexual reproductive structures. In bryophytes, dioecy and sex ratio bias is especially common, making them effective organisms for studying sex ratio bias. Given the frequency of female bias in bryophytes, the hypothesis was that females will show a higher competitive ability relative to males. The experiment was conducted in greenhouse conditions at several densities and proportions over the course of seven months. As individuals grew and formed clumps, identities were tracked, and growth measurements were made using photographs and computer imaging software. Both sexes grew on average 41% more with the opposite sex relative to their single-sex cultures. A model predicting future sex ratios showed coexistence between the sexes and predicted a male biased sex ratio of 3.2 males to 1 female. A trade-off was observed for males where single-sex cultures contained more asexual structures than mixed-sex cultures and the reverse for growth rate. Higher levels of asexual reproduction in males in single-sex cultures might be selected for to increase male dispersal for a higher probability of encountering females. This pattern was not found for females. The overyielding results suggest an interaction effect may exist due to niche differentiation between the sexes. In addition, the results suggest that in some dioecious plants a change in sex ratio can occur before differences in their allocation to sexual reproduction.

Les prairies permanentes représentent l’une des principales formes d’utilisation des terres en Europe. En France, elles représentent près de 40% de la Surface Agricole Utile (SAU) et restent le support de nombreux systèmes d’élevage. Les scénarios actuels de changement climatique prévoient tous un accroissement de la variabilité climatique qui pourrait se traduire par un accroissement des évènements extrêmes, notamment en ce qui concerne les épisodes de sécheresses. Ces événements extrêmes sont de nature à altérer la structure et le fonctionnement des prairies et en conséquence la capacité de ces écosystèmes à rendre des services agricoles, écologiques et environnementaux. Une meilleure compréhension des mécanismes de résistance et de résilience des communautés végétales prairiales face à l’occurrence de sécheresses extrêmes s’avère donc cruciale pour évaluer la stabilité du fonctionnement de l’écosystème prairial et notamment du service de production qu'il supporte. Cette thèse a pour objectif principal d’évaluer la capacité des prairies permanentes à maintenir leur fonction de production face à un événement de sécheresse extrême. Elle s’appuie sur une démarche expérimentale en mésocosmes sous conditions semi-contrôlées permettant de caractériser le rôle de la composition et de la structure des communautés végétales prairiales dans la résistance et la résilience à la sécheresse des prairies. Pour évaluer l’importance de la composition et de la diversité fonctionnelle dans la réponse des communautés végétales, des monocultures et des mélanges associant graminées et une légumineuse ont été sélectionnés. L’originalité de ce travail réside dans une approche d’écologie fonctionnelle permettant d’appréhender l’importance des interactions entre espèces dans la réponse de la communauté via l’analyse de traits morphologiques et physiologiques mesurés conjointement dans les compartiments aérien et souterrain de la végétation. Cette étude a mis en évidence le rôle prépondérant du Trèfle blanc (Trifolium repens) dans la modulation de l’impact d’un événement de sécheresse extrême sur la production de biomasse de mélanges prairiaux. Si aucun effet tampon de la composition et de la structure de la communauté n’a été mis en évidence durant la sécheresse, la présence et l’abondance de Trèfle blanc a été reliée à une meilleure récupération à long terme de la production de biomasse des mélanges. Après l’extrême, une surproduction des mélanges associant graminée et Trèfle blanc a ainsi été attribuée à des mécanismes de facilitation azotée et de complémentarité verticale de niches au niveau racinaire pour l’acquisition de l’eau. Nous avons ainsi souligné l’importance de la croissance des racines profondes pour la mise en place de complémentarités fonctionnelles entre espèces et donc pour le maintien de la production sous sécheresse extrême. L’événement de sécheresse affecte durablement le comportement des racines et de ce fait les mécanismes sous-jacents de l’effet bénéfique du Trèfle blanc sur la production aérienne. Par ailleurs, notre étude a mis en évidence le rôle majeur des interactions hétérospécifiques dans l’expression de traits intraspécifiques liés aux stratégies d’acquisition des ressources et par conséquent dans la régulation de la performance des espèces sous sécheresse extrême. Nous montrons que la structure et la composition fonctionnelle de la communauté végétale apparaît essentielle dans la régulation de la réponse à un événement de sécheresse extrême. Dans notre étude, la récupération de la production de biomasse aérienne après sécheresse extrême repose essentiellement sur la présence de Trèfle blanc et la mise en place d’interactions positives entre espèces. La prise en compte de la modulation par les interactions interspécifiques du comportement des espèces en mélanges s’avère nécessaire dans la prévision de la production des prairies sous climat futur. (...) / Permanent grasslands are one of the main forms of land use in Europe. In France, permanent grasslands represent around 40% of Utilised Agricultural Land and support many animal-based farming systems. Current climate change scenarios predict increased climate variability that could result in an increase in extreme events such as drought episodes. These extreme events can significantly alter the structure and functioning of grasslands, with implications for their capacity to provide agricultural, ecological and environmental services. Improved understanding of the resistance and resilience mechanisms of grassland plant communities exposed to extreme drought is therefore critical in order to assess the stability of grassland functions, in particular that of biomass production. This thesis aims at assessing the capacity of permanent grasslands to sustain their production when exposed to an extreme drought event. The study uses an experimental approach and semi-controlled conditions to investigate the role of community composition and functional diversity for drought resistance and resilience in grassland mixtures. Monocultures and mixtures combining grasses and legumes were selected to test the importance of composition and functional diversity for plant community responses during drought and up to a year after the end of water stress. The originality of this work is the use of a functional ecological approach to assess the impacts of species’ interactions via measurements of plant morphological and physiological traits both above- and below-ground. This study demonstrates the importance of white clover (Trifolium repens) in modulating the impact of extreme droughts on biomass production of grassland mixtures. Community composition did not have a buffering effect on biomass production during the drought event but the presence and abundance of white clover in mixtures was linked to higher recovery of biomass production in the long-term. After the extreme drought event, overyielding in mixtures combining grass and white clover could be attributed to mechanisms of nitrogen facilitation and vertical root complementarity for water uptake. Results also show the importance of deep root growth for the establishment of functional complementarity between species and the maintenance of biomass production under extreme drought. Drought had lasting effects on the root system patterns and thus on the underlying mechanisms of the positive effect of white clover on aboveground production. Moreover, this work highlights the key role of heterospecific interactions in the expression of intraspecific traits related to resource acquisition strategies. Heterospecific interactions are thus an important driver of the species performance under extreme drought.We show that the structure and functional composition of the plant community in simplified mixtures are critical for modulating grassland responses to an extreme drought event. In our study, the recovery of aboveground biomass production is mainly related to the presence of white clover and the establishment of positive interactions between species. Changes of species behaviour in response to heterospecific interactions should be taken into account to better predict grassland biomass production under future climatic conditions. From an applied perspective, accounting for the key role of white clover and the inherent variability or neighbour-induced plasticity of intraspecific traits appears important to adapt mixtures of species or varieties to more intense and frequent drought events.

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Diverse native seed mixtures have many benefits for prairie restoration or seeded pastures. In natural grasslands, species naturally coexist with hundreds of other species in complex communities. Commercial seed mixtures rarely contain more than a small number of species, often with haphazard ratios of the component species. Thus there is no natural template for combining selected species into an optimally productive community and there is limited knowledge on how to compose a suitable species mixture. Identifying which features of a community drive increased productivity may aid in screening species and community compositions, leading to mixtures that are more specifically designed to be stable, and highly productive for the region. There is renewed interest native species as they have the potential to provide non-invasive, productive, and drought resistant rangelands that may prove more sustainable. Seven species with high agronomic potential and a broad native geographic distribution were selected for testing including: nodding brome [Bromus anomalus (Coult.)], blue bunch wheatgrass [Pseudoregneria spicata (Pursh)], western wheatgrass [Pascopyrum smithii (Rydb.)], side oats grama [Bouteloua curtipendula (Michx.)], little blue stem [Schizachyrium scoparium (Michx.)], purple prairie clover [Dalea purpurea (Vent.)], and white prairie clover [Dalea candida (Willd.)]. The early productivity and nutritional quality of these species was determined in simple mixtures in two field sites: Saskatoon and Swift Current. In the field sites the mixtures included all seven monocultures, 21 two-species mixtures and a mixture with all species. Productivity may be driven by the species richness, functional group richness, and species evenness of the community, the abundance and occurrence of particular species or functional groups, and average plant trait values within the community. Therefore, identifying the features of a community that drive increased productivity and applying them as predictive tools may aid in screening species and community compositions. Many complex mixtures of the species were planted in greenhouse experiments to determine the strongest drivers of productivity for communities of these species. The experimental approach was validated in a confirmatory experiment where optimum communities were tested. These results did not differ under a moderate drought treatment. Results were generally consistent between field and greenhouse studies. Western wheatgrass (WWG) had the highest overall plant density and the strongest effect on the forage yield of the mixtures and communities. In the field study, productivity and crude protein content were not reduced when other species were also included with WWG in the mixture. Dalea spp. did not establish as well as the other species, but had the highest crude protein concentrations. The strongest predictors of productivity were the presence and abundance of perennial C3 grasses. Increases in species richness, functional group richness, and the presence of C3s (more specifically western wheatgrass) also increased productivity, likely because of the high early relative growth rate and strong competitive ability of western wheatgrass. Overall, communities screened in the greenhouse reflected early establishment field results. The systematic approach for evaluating communities can be modified to consider enhancing other ecological functions in addition to high productivity, in other regions.

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