Sommaire : Comprendre le fonctionnement des écosystèmes est un enjeu crucial, en particulier dans un contexte de changements globaux. Afin de mieux prédire les processus écosystémiques, j’ai testé la précision et les limites des hypothèses du biomass-ratio de Grime (HBMR) et de l’annulation idiosyncratique (HAI), cette dernière étant une hypothèse originale de cette thèse. Pour cela, j’ai appliqué le principe du biomass-ratio aux traits fonctionnels, en employant la méthode des traits agrégés en communauté, pour estimer la réponse globale des espèces en mélange. La décomposition des litières plurispécifiques constitue un bon modèle biologique, pour lequel je me suis posé les questions suivantes : (1) est-ce que l’HBMR prédit bien les taux de décomposition en mélanges plurispécifiques ? ; (2) est-ce que le degré de variabilité de ces taux diminue pour des raisons biologiques avec l’augmentation de la richesse spécifique (RS) des mélanges (HAI) ? ; (3) est-ce que la variabilité des taux entre mélanges diminue quand les conditions abiotiques du site deviennent plus limitantes ? ; (4) considérant que les mélanges plus contrastés fonctionnellement sont susceptibles de développer plus d’interactions, est-ce que la déviation à la prédiction augmente avec la dispersion fonctionnelle des mélanges (« FDis », Laliberté & Legendre 2010) ? Cette thèse inclut deux expériences de décomposition en sachets à litières : (1) à Sherbrooke (QC, Canada) avec des microcosmes, impliquant des litières de six espèces d’arbres, décomposant seules et en mélanges et (2) sur trois sites au climat contrasté dans la région de Clermont-Ferrand (France) avec des litières de quatre espèces d’herbacées, décomposant seules et en mélanges. Les résultats montrent des déviations positives et négatives par rapport aux taux prédits, mais l’HBMR décrit bien la réponse moyenne des litières plurispécifiques. Bien que l’HAI ait été rejetée, les résultats montrent une convergence des taux observés vers les taux prédits quand (1) la RS des mélanges augmente, (2) l’échelle spatiale augmente et (3) le climat est plus limitant pour la décomposition. Enfin, malgré des corrélations entre FDis et interactions entre espèces dans les litières, cette relation n’est pas généralisable et l’hypothèse de corrélation positive entre FDis et déviation à l’HBMR a été rejetée. // Abstract : Understanding ecosystem functioning is a key goal in ecology, especially in the context of global changes. To better predict ecosystem processes, I tested the accuracy and the limits of Grime’s biomass-ratio (BMRH) hypothesis and a novel idiosyncratic annulment (IAH) hypothesis. I applied the biomass-ratio to functional traits, using the community-weighted means (CWM) to estimate the global response of species in mixtures. I studied the decomposition of litter species mixtures as a biological model and asked the following questions : (1) does the BMRH predict well the decomposition rates of mixed species litters? ; (2) does the degree of variability of these rates decrease with increasing species richness (SR) beyond that expected from purely mathematical causes (IAH)? ; (3) does the variability of rates between mixtures decrease with less favourable abiotic conditions for decomposition? ; (4) as more functionally contrasted mixtures are expected to develop more interactions, does the deviation from prediction increase with increasing functional dispersion in mixtures (« FDis », Laliberté & Legendre 2010)? This study involves two decomposition experiments using litterbags: (1) at Sherbrooke (QC, Canada), in microcosms, involving litters from six tree species, decomposed alone and in mixtures and (2) in three climatically contrasted sites in the region of Clermont-Ferrand (France) with litters from four herbaceous species, decomposed alone and in mixtures. Despite both positive and negative deviations from expectation occurring at all levels of SR, the BMRH well described the average response of mixed species litters. Although I rejected the IAH, the results showed a convergence to the predicted values based on CWM with (1) increasing the SR in mixtures, (2) increasing the spatial scale of the study and (3) a less favourable climate to decomposition. Finally, although there was a correlation between litter interactions and functional divergence, this relationship was not generalizable and I rejected the hypothesis of a positive correlation between FDis and the deviations from BMRH.

Despite the importance of a high white clover (Trifolium repens) content in temperate pastoral systems in terms of livestock performance and nitrogen fixation, the proportion of white clover in grass-clover pastures is often low (<20%). This thesis examined in two experiments whether the white clover content of pastures could be improved by sowing white clover with alternative grass species to diploid perennial ryegrass (Lolium perenne L.). In a pasture experiment, DM production, pasture composition and morphology of grass-clover mixtures was measured over the establishment year (January 2007 to January 2008) where white clover was sown in fine mixtures with diploid perennial ryegrass, tetraploid perennial ryegrass, timothy (Phleum pratense L.) and cocksfoot (Dactylis glomerata L.). Pastures were irrigated and rotationally grazed with on-off grazing with Coopworth ewe hoggets. Total annual DM production of pasture was more than 20% higher in tetraploid (12521 kg DM ha⁻¹) and diploid (11733 kg DM ha⁻¹) perennial ryegrass than timothy (9751 kg DM ha⁻¹) and cocksfoot (9654 kg DM ha⁻¹). However, timothy (5936 kg DM ha⁻¹) and cocksfoot (5311 kg DM ha⁻¹) had more than four times higher white clover annual DM production than tetraploid (1310 kg DM ha⁻¹) and diploid (818 kg DM ha⁻¹) ryegrass. Pasture growth rate at the first three harvests in autumn was significantly greater in tetraploid and diploid ryegrass than timothy and cocksfoot. Timothy and cocksfoot had a higher proportion of white clover than tetraploid and diploid perennial ryegrass throughout the entire year. This was due to more and larger white clover plants in timothy and cocksfoot plots. In a grazing preference experiment, the partial preference of sheep for white clover offered in combination with the same grass species as in the pasture experiment was measured in five grazing tests in May, September, October, November and December 2007. Pastures were sown in January 2007. Paired plots (grass and clover both 4.2 m x 10 m) were grazed by three Coopworth ewe hoggets between 9am and 5pm, and preference was recorded by decline in pasture mass and visual scan sampling for grazing time. Grazing preference for clover was generally low throughout these tests (e.g. average apparent DM intake from clover = 47%; average grazing time from clover = 44%). Several explanations are proposed for this low preference including a high N content and intake rate of the grass relative to the clover. No significant differences were found among the grass treatments in total grass grazing time, total clover grazing time, ruminating time, the proportion of grazing time on clover, selective coefficient for clover and DM intake percentage from clover at any date. There was no significant change in overall sward surface height (SSH) decline among grass treatments throughout all the tests except December 2007 when the overall SSH decline for cocksfoot was significantly lower than the other species. The study indicated that the rapid growth rate of perennial ryegrass in the early phase of pasture establishment, rather than differences in partial preference, was the key factor limiting white clover content in the mixed swards relative to cocksfoot and timothy pastures. It is concluded that high clover-containing pastures capable of delivering high per head performance can be established through the use of slow establishing pasture species such as timothy and cocksfoot.

Although New Zealand is fortunate in having a climate mild enough to allow some growth of pasture in winter even in the extreme south of the South Island (Duffy, 1971), growth is nevertheless considerably less than in spring and summer. O’Connor et al, (1968) point out that at Lincoln, Canterbury, winter production from a New Zealand Certified Grasslands Ruanui Perennial ryegrass (Lolium perenne c.v.) and New Zealand Certified Grasslands Huia white clover (Trifolium repens c.v.) sward is at best only 8% of the mean total annual dry matter (D.M.) production. A similar seasonal pattern is shown by irrigated pastures (Rickard, 1968), as well as pastures in the milder North Island of New Zealand (O’Connor and Vartha, pers comm.). Stocking of grazing animals is normally related to the growth rate of pastures in the spring, with conserved hay or silage, plus specially grown greenfeed or root crops being fed if necessary to offset the winter feed deficit. However on hill country too steep for tractor cultivation, cropping and conservation are not possible. Animals have to rely upon in situ grazing of native or dominant browntop (Agrostis tenuis) swards which may not have been improved by the aerial introduction of clovers. In an unimproved state, the quality and quantity of the herbage grown on such areas are low, Molloy (1966) have discussed clover introduction into native swards and the notably beneficial result this has on stock thrift, particularly during the winter period. Considerably better growth rates of stock grazing legumes rather than grasses have been noted by many authors including Ulyatt, (1971), and McLean et al (1962); this superiority being due largely to the greater digestible organic matter intake and higher ratio of soluble to insoluble carbohydrate associated with the herbage of legumes (Ulyatt, 1971). Consequently the presence of clover, and the winter activity of this, have an important influence on the productivity of steep hill country during winter. An increase of winter activity is also desirable, but not essential, in clover associated with flat or gently rolling pastures. The main pasture legume used in New Zealand has, in the past, been the white clover cultivar Huia. Whilst since 1945 selection and breeding of ryegrasses has brought about a very considerable improvement in the winter or cool season activity of these (Corkill, 1966), no legume cultivar having an increased level of cool-season activity has been released to commerce in New Zealand over the same period to date. Breeding for increased winter growth has been an objective of the Grasslands Division Plant Breeding Section for many years (Barclay, 1960). Since 1957, breeding and selection work has been proceeding with the objective of increasing the winter growth of the New Zealand Certified Grasslands white clover cultivar – Huia (henceforth to be referred to as Huia) without sacrificing the moderately good summer growth of this strain, (Barclay, 1969). Seed of a promising cultivar selected during the course of this work – New Zealand Grasslands 4700 white clover, (henceforth to be referred to as 4700) became available for testing in 1967. The investigation to be reported here aimed to measure the cool-season activity of 4700 by comparison with that of Huia, at the same time elucidating if possible the factors controlling this growth.

The annual and seasonal water use efficiency of six pasture combinations were calculated from the ‘MaxClover’ Grazing Experiment at Lincoln University. Pastures have been established for six years and are grazed by best management practices for each combination. Measurements for this study are from individual plots of four replicates of ryegrass (RG)/white clover (Wc), cocksfoot (CF)/Wc; CF/balansa (Bal) clover; CF/Caucasian (Cc) clover; CF/subterranean (Sub) clover or lucerne. Water extraction measurements showed soils for all dryland pastures had a similar plant available water content of 280±19.8 mm. Dry matter measurements of yield, botanical composition and herbage quality were assessed from 1 July 2008 until 30 June 2009. Lucerne had the highest annual yield of 14260 kg DM/ha/y followed by the CF/Sub at 9390 kg DM/ha/y and the other grass based pastures at ≤ 6900 kg DM/ha/y. All pastures used about 670±24.4 mm/y of water for growth. Lucerne had the highest annual water use efficiency (WUE) of 21 kg DM/ha/mm/y of water used (total yield/total WU). The WUE of CF/Sub was the second highest at 15 kg DM/ha/mm/y, and the lowest was CF/Wc at 9 kg DM/ha/mm/y. The CF/Sub pastures had the highest total legume content of all grass based pastures at 21% and as a consequence had the highest annual nitrogen yield of 190 kg N/ha. This was lower than the monoculture of lucerne (470 kg N/ha). Ryegrass/white clover had the highest total weed component in all pastures of 61%. For dryland farmers spring is vital for animal production when soil temperatures are rising and moisture levels are high. The water use efficiency at this time is important to maximize pasture production. In spring lucerne produced 8730 kg DM/ha, which was the highest dry matter yield of all pastures. The CF/Sub produced the second highest yield of 6100 kg/DM/ha. When calculated against thermal time, CF/Sub grew 5.9 kg DM/ºCd compared with lucerne at 4.9 kg DM/ºCd. The higher DM yield from lucerne was from an extra 400 ºCd of growth. The highest seasonal WUE of all pastures occurred in the spring growing period. Linear regressions forced through the origin, showed lucerne (1/7/08-4/12/08) had a WUE of 30 kg DM/ha/mm (R2=0.98). Of the grass based pastures, CF/Sub produced 18 kg DM/ha/mm (R2=0.98) from 1/7 to 10/11/08 from 270 mm of water used. The lowest spring WUE was 13.5 kg DM/ha/mm by CF/Bal pastures which was comparable to the 14.3±1.42 kg DM/ha/mm WUE of CF/Wc, CF/Cc and RG/Wc pastures. During the spring, CF/Sub clover had the highest spring legume component of the grass based pastures at 42% and produced 120 kg N/ha. This was lower than the 288 kg N/ha from the monoculture of lucerne. Sub clover was the most successful clover which persisted with the cocksfoot. Based on the results from this study dryland farmers should be encouraged to maximize the potential of lucerne on farm, use cocksfoot as the main grass species for persistence, rather than perennial ryegrass, and use subterranean clover as the main legume species in cocksfoot based pastures. By increasing the proportion of legume grown the water use efficiency of a pasture can be improved. When pastures are nitrogen deficient the use of inorganic nitrogen may also improve pasture yields particularly in spring.

Sommaire : Comprendre le fonctionnement des e??cosyste??mes est un enjeu crucial, en particulier dans un contexte de changements globaux. Afin de mieux pre??dire les processus e??cosyste??miques, j???ai teste?? la pre??cision et les limites des hypothe??ses du biomass-ratio de Grime (HBMR) et de l???annulation idiosyncratique (HAI), cette dernie??re e??tant une hypothe??se originale de cette the??se. Pour cela, j???ai applique?? le principe du biomass-ratio aux traits fonctionnels, en employant la me??thode des traits agre??ge??s en communaute??, pour estimer la re??ponse globale des espe??ces en me??lange. La de??composition des litie??res plurispe??cifiques constitue un bon mode??le biologique, pour lequel je me suis pose?? les questions suivantes : (1) est-ce que l???HBMR pre??dit bien les taux de de??composition en me??langes plurispe??cifiques ? ; (2) est-ce que le degre?? de variabilite?? de ces taux diminue pour des raisons biologiques avec l???augmentation de la richesse spe??cifique (RS) des me??langes (HAI) ? ; (3) est-ce que la variabilite?? des taux entre me??langes diminue quand les conditions abiotiques du site deviennent plus limitantes ? ; (4) conside??rant que les me??langes plus contraste??s fonctionnellement sont susceptibles de de??velopper plus d???interactions, est-ce que la de??viation a?? la pre??diction augmente avec la dispersion fonctionnelle des me??langes (?? FDis ??, Laliberte?? & Legendre 2010) ? Cette the??se inclut deux expe??riences de de??composition en sachets a?? litie??res : (1) a?? Sherbrooke (QC, Canada) avec des microcosmes, impliquant des litie??res de six espe??ces d???arbres, de??composant seules et en me??langes et (2) sur trois sites au climat contraste?? dans la re??gion de Clermont-Ferrand (France) avec des litie??res de quatre espe??ces d???herbace??es, de??composant seules et en me??langes. Les re??sultats montrent des de??viations positives et ne??gatives par rapport aux taux pre??dits, mais l???HBMR de??crit bien la re??ponse moyenne des litie??res plurispe??cifiques. Bien que l???HAI ait e??te?? rejete??e, les re??sultats montrent une convergence des taux observe??s vers les taux pre??dits quand (1) la RS des me??langes augmente, (2) l???e??chelle spatiale augmente et (3) le climat est plus limitant pour la de??composition. Enfin, malgre?? des corre??lations entre FDis et interactions entre espe??ces dans les litie??res, cette relation n???est pas ge??ne??ralisable et l???hypothe??se de corre??lation positive entre FDis et de??viation a?? l???HBMR a e??te?? rejete??e. // Abstract : Understanding ecosystem functioning is a key goal in ecology, especially in the context of global changes. To better predict ecosystem processes, I tested the accuracy and the limits of Grime???s biomass-ratio (BMRH) hypothesis and a novel idiosyncratic annulment (IAH) hypothesis. I applied the biomass-ratio to functional traits, using the community-weighted means (CWM) to estimate the global response of species in mixtures. I studied the decomposition of litter species mixtures as a biological model and asked the following questions : (1) does the BMRH predict well the decomposition rates of mixed species litters? ; (2) does the degree of variability of these rates decrease with increasing species richness (SR) beyond that expected from purely mathematical causes (IAH)? ; (3) does the variability of rates between mixtures decrease with less favourable abiotic conditions for decomposition? ; (4) as more functionally contrasted mixtures are expected to develop more interactions, does the deviation from prediction increase with increasing functional dispersion in mixtures (?? FDis ??, Lalibert?? & Legendre 2010)? This study involves two decomposition experiments using litterbags: (1) at Sherbrooke (QC, Canada), in microcosms, involving litters from six tree species, decomposed alone and in mixtures and (2) in three climatically contrasted sites in the region of Clermont-Ferrand (France) with litters from four herbaceous species, decomposed alone and in mixtures. Despite both positive and negative deviations from expectation occurring at all levels of SR, the BMRH well described the average response of mixed species litters. Although I rejected the IAH, the results showed a convergence to the predicted values based on CWM with (1) increasing the SR in mixtures, (2) increasing the spatial scale of the study and (3) a less favourable climate to decomposition. Finally, although there was a correlation between litter interactions and functional divergence, this relationship was not generalizable and I rejected the hypothesis of a positive correlation between FDis and the deviations from BMRH.

Comprendre le fonctionnement des écosystèmes est un enjeu crucial, en particulier dans un contexte de changements globaux. Afin de mieux prédire les processus écosystémiques, j'ai testé la précision et les limites des hypothèses du biomass-ratio de Grime (HBMR) et de l'annulation idiosyncratique (HAI), cette dernière étant une hypothèse originale de cette thèse. Pour cela, j'ai appliqué le principe du biomass-ratio aux traits fonctionnels, en employant la méthode des traits agrégés en communauté, pour estimer la réponse globale des espèces en mélange. La décomposition des litières plurispécifiques constitue un bon modèle biologique, pour lequel je me suis posé les questions suivantes : (1) est-ce que l'HBMR prédit bien les taux de décomposition en mélanges plurispécifiques ? ; (2) est-ce que le degré de variabilité de ces taux diminue pour des raisons biologiques avec l'augmentation de la richesse spécifique (RS) des mélanges (HAI) ? ; (3) est-ce que la variabilité des taux entre mélanges diminue quand les conditions abiotiques du site deviennent plus limitantes ? ; (4)considérant que les mélanges plus contrastés fonctionnellement sont susceptibles de développer plus d'interactions, est-ce que la déviation à la prédiction augmente avec la dispersion fonctionnelle des mélanges(" FDis ", La liberté & Legendre 2010) ? Cette thèse inclut deux expériences de décomposition en sachets à litières : (1) à Sherbrooke (QC, Canada) avec des microcosmes, impliquant des litières de six espèces d'arbres, décomposant seules et en mélanges et (2) sur trois sites au climat contrasté dans la région de Clermont-Ferrand (France) avec des litières de quatre espèces d'herbacées, décomposant seules et en mélanges. Les résultats montrent des déviations positives et négatives par rapport aux taux prédits, mais l'HBMR décrit bien la réponse moyenne des litières plurispécifiques. Bien que l'HAI ait été rejetée, les résultats montrent une convergence des taux observés vers les taux prédits quand (1) la RS des mélanges augmente, (2)l'échelle spatiale augmente et (3) le climat est plus limitant pour la décomposition. Enfin, malgré des corrélations entre FDis et interactions entre espèces dans les litières, cette relation n'est pas généralisable et l'hypothèse de corrélation positive entre FDis et déviation à l'HBMR a été rejetée.