• The Agrodiversity Experiment: three years of data from a multisite study in intensively managed grasslands

      Kirwan, Laura; Connolly, John; Brophy, C.; Baadshaug, Ole; Belanger, Gilles; Black, Alistair D; Camus, Tim; Collins, Rosemary; Cop, Jure; Delgado, Ignacio; et al. (Ecological Society of America, 11/06/2014)
      Intensively managed grasslands are globally prominent ecosystems. We investigated whether experimental increases in plant diversity in intensively managed grassland communities can increase their resource use efficiency. This work consisted of a coordinated, continental-scale 33-site experiment. The core design was 30 plots, representing 15 grassland communities at two seeding densities. The 15 communities were comprised of four monocultures (two grasses and two legumes) and 11 four-species mixtures that varied in the relative abundance of the four species at sowing. There were 1028 plots in the core experiment, with another 572 plots sown for additional treatments. Sites agreed a protocol and employed the same experimental methods with certain plot management factors, such as seeding rates and number of cuts, determined by local practice. The four species used at a site depended on geographical location, but the species were chosen according to four functional traits: a fast-establishing grass, a slow-establishing persistent grass, a fast-establishing legume, and a slow-establishing persistent legume. As the objective was to maximize yield for intensive grassland production, the species chosen were all high-yielding agronomic species. The data set contains species-specific biomass measurements (yield per species and of weeds) for all harvests for up to four years at 33 sites. Samples of harvested vegetation were also analyzed for forage quality at 26 sites. Analyses showed that the yield of the mixtures exceeded that of the average monoculture in >97% of comparisons. Mixture biomass also exceeded that of the best monoculture (transgressive overyielding) at about 60% of sites. There was also a positive relationship between the diversity of the communities and aboveground biomass that was consistent across sites and persisted for three years. Weed invasion in mixtures was very much less than that in monocultures. These data should be of interest to ecologists studying relationships between diversity and ecosystem function and to agronomists interested in sustainable intensification. The large spatial scale of the sites provides opportunity for analyses across spatial (and temporal) scales. The database can also complement existing databases and meta-analyses on biodiversity–ecosystem function relationships in natural communities by focusing on those same relationships within intensively managed agricultural grasslands.
    • Ecosystem function enhanced by combining four functional types of plant species in intensively-managed grassland mixtures: a three-year continental-scale field experiment

      Finn, John; Kirwan, Laura; Connolly, John; Sebastia, Maria Teresa; Helgadottir, Aslaug; Baadshaug, Ole; Belanger, Gilles; Black, Alistair D; Brophy, C.; Collins, Rosemary; et al. (Wiley-Blackwell, 22/02/2013)
      1. A co-ordinated continental-scale field experiment across 31 sites was used to compare the biomass yield of monocultures and four-species mixtures associated with intensively managed agricultural grassland systems. To increase complementarity in resource use, each of the four species in the experimental design represented a distinct functional type derived from two levels of each of two functional traits, nitrogen acquisition (N2-fixing legume or non-fixing grass) crossed with temporal development (fast-establishing or temporally persistent). Relative abundances of the four functional types in mixtures were systematically varied at sowing to vary the evenness of the same four species in mixture communities at each site, and sown at two levels of seed density. 2. Across multiple years, the total yield (including weed biomass) of the mixtures exceeded that of the average monoculture in >97% of comparisons. It also exceeded that of the best monoculture (transgressive overyielding) in about 60% of sites, with a mean yield ratio of mixture to best-performing monoculture of 1.07 across all sites. Analyses based on yield of sown species only (excluding weed biomass) demonstrated considerably greater transgressive overyielding (significant at about 70% of sites, ratio of mixture to best-performing monoculture = 1.18). 3. Mixtures maintained a resistance to weed invasion over at least three years. In mixtures, median values indicate <4% of weed biomass in total yield, whereas the median percentage of weeds in monocultures increased from 15% in year 1 to 32% in year 3. 4. Within each year, there was a highly significant relationship (P<0.0001) between sward evenness and the diversity effect (excess of mixture performance over that predicted from the monoculture performances of component species). At lower evenness values, increases in community evenness resulted in an increased diversity effect, but the diversity effect was not significantly different from the maximum diversity effect across a wide range of higher evenness values. The latter indicates the robustness of the diversity effect to changes in species’ relative abundances. 5. Across sites with three complete years of data (24 of the 31 sites), the effect of interactions between the fast-establishing and temporal persistent trait levels of temporal development was highly significant and comparable in magnitude to effects of interactions between N2-fixing and non-fixing trait levels of nitrogen acquisition. 6. Synthesis and applications. The design of grassland mixtures is relevant to farm-level strategies to achieve sustainable intensification. Experimental evidence indicated significant yield benefits of four-species agronomic mixtures which yielded more than the highest-yielding monoculture at most sites. The results are relevant for agricultural practice, and show how grassland mixtures can be designed to improve resource complementarity, increase yields and reduce weed invasion. The yield benefits were robust to considerable changes in the relative proportions of the four species, which is extremely useful for practical management of grassland swards.
    • Effects of multi-species swards on dry matter production and the incidence of unsown species at three Irish sites

      Connolly, John; Finn, John; Black, Alistair D; Kirwan, Laura; Brophy, C.; Luscher, A.; European Union; Science Foundation Ireland; COST Action 852; 09/RFP/EOB2546 (Teagasc (Agriculture and Food Development Authority), Ireland, 2009)
      Recent ecological research provides evidence that an increased number of plant species in natural grasslands is associated with increased biomass productivity, and provides a wide range of other ecosystem benefits. This suggests that increases in species diversity in agricultural ecosystems may similarly lead to increased benefits. The work reported below was part of the COST 852 Agrodiversity experiment, carried out at 34 sites across Europe. In Ireland, the effects of four-species grass-clover mixtures on herbage production, species persistence and unsown species suppression at three sites over multiple years, were investigated under growing conditions that were intensive relative to unfertilised natural grassland systems. The design included a range of four-species mixtures and monocultures of perennial ryegrass, timothy, cocksfoot, white clover, red clover and Caucasian clover. Several harvests were taken at each site for two or three years. Species diversity had a strong, persistent and positive effect on overall yield and the yield of sown species, and enhanced resistance to the growth of unsown species. Mixtures generally yielded well when compared with the best monoculture, and sometimes out yielded it. These effects on total yield declined over time but were still important at the end of the experiments. The diversity effects on sown species yield and on resistance to unsown species increased with time. Diversity effects were robust to changes in species composition, and persisted for the duration of the experiments across mixtures and over time. Virtually every mixture had a higher yield, and suppressed unsown species better, than monocultures of perennial ryegrass. These patterns were broadly consistent across sites. The persistence of species varied widely and was not consistent across sites.
    • Nitrogen yield advantage from grass-legume mixtures is robust over a wide range of legume proportions and environmental conditions

      Suter, Matthias; Connolly, John; Finn, John; Loges, R.; Kirwan, Laura; Sebastia, Maria Teresa; Luscher, A.; European Union (Wiley, 28/01/2015)
      Current challenges to global food security require sustainable intensification of agriculture through initiatives that include more efficient use of nitrogen (N), increased protein self-sufficiency through home-grown crops, and reduced N losses to the environment. Such challenges were addressed in a continental-scale field experiment conducted over three years, in which the amount of total nitrogen yield (Ntot) and the gain of N yield in mixtures as compared to grass monocultures (Ngainmix) was quantified from four-species grass-legume stands with greatly varying legume proportions. Stands consisted of monocultures and mixtures of two N2 fixing legumes and two non-fixing grasses.The amount of Ntot of mixtures was significantly greater (P ≤ 0.05) than that of grass monocultures at the majority of evaluated sites in all three years. Ntot and thus Ngainmix increased with increasing legume proportion up to one third of legumes. With higher legume percentages, Ntot and Ngainmix did not continue to increase. Thus, across sites and years, mixtures with one third proportion of legumes attained ~95% of the maximum Ntot acquired by any stand and had 57% higher Ntot than grass monocultures.Realized legume proportion in stands and the relative N gain in mixture (Ngainmix/Ntot in mixture) were most severely impaired by minimum site temperature (R = 0.70, P = 0.003 for legume proportion; R = 0.64, P = 0.010 for Ngainmix/Ntot in mixture). Nevertheless, the relative N gain in mixture was not correlated to site productivity (P = 0.500), suggesting that, within climatic restrictions, balanced grass-legume mixtures can benefit from comparable relative gains in N yield across largely differing productivity levels.We conclude that the use of grass-legume mixtures can substantially contribute to resource-efficient agricultural grassland systems over a wide range of productivity levels, implying important savings in N fertilizers and thus greenhouse gas emissions and a considerable potential for climate change mitigation.
    • Yield of temperate forage grassland species is either largely resistant or resilient to experimental summer drought

      Hofer, Daniel; Suter, Matthias; Haughey, Eamon; Finn, John; Hoekstra, Nyncke J.; Buchmann, Nina; Luscher, A.; European Union; 266018 (Wiley, 17/06/2016)
      1.Due to climate change, an increasing frequency and severity of drought events are expected to impair grassland productivity, particularly of intensively managed temperate grasslands. 2.To assess drought impacts, a common field experiment to manipulate precipitation was set up at three sites (two Swiss and one Irish) using monocultures and mixtures with two and four key forage species. Species differed in their functional traits: a shallow-rooted non-legume (Lolium perenne L.), a deep-rooted non-legume (Cichorium intybus L.), a shallow-rooted legume (Trifolium repens L.) and a deep-rooted legume (Trifolium pratense L.). A 9-week summer drought was simulated, and soil water status, above-ground biomass yield and plant nitrogen (N) limitation were compared to a rainfed control. 3.Based on soil water measurements, the drought induced severe stress at both Swiss sites and extreme stress at the Irish site. Under severe stress, the legumes were more drought resistant and showed an average change in above-ground biomass (CAB, compared to rainfed control) of only −8% and −24% (for the two Swiss sites), while the non-legumes had an average CAB of −51% and −68%. The lower resistance of non-legumes coincided with an apparent limitation of plant N, which further increased under drought. Under extreme drought (Irish site), growth nearly ceased with an average CAB of −85%. 4.During a 6-week post-drought period with adequate water supply (Swiss sites), formerly drought-stressed species were highly resilient and either attained (legumes) or clearly outperformed (non-legumes) the yield level of the rainfed controls. This outperformance coincided with post-drought reductions in N limitation in formerly drought-stressed species. As a result, aggregated over the drought and the post-drought periods, a negative drought impact was found only for the shallow-rooted L. perenne at one of the severely stressed sites. 5.Significant overyielding by multispecies mixtures was evident under rainfed control conditions (+38% across all three sites, P < 0·05) and was equally apparent under severe drought (+50%, P < 0·05). This overyielding was greatest in mixtures with approximately equal species proportions and was sufficiently large that drought-stressed mixtures at least attained the same yield as the average of the rainfed monocultures. Under extreme drought, growth almost ceased in monocultures and mixtures. 6.Synthesis and applications. Yields of selected species of intensively managed temperate grasslands are either resistant to a single severe drought or are highly resilient as soon as soil moisture levels recover after the drought event. However, these forage species seem unable to cope with an extreme drought event. Combining species in mixtures can compensate for yield reductions caused by severe drought and it offers a practical management tool to adapt forage production to climate change.