Integrating design and ecological theory to achieve adaptive diverse pastures.

Increasing plant diversity is often suggested as a way of overcoming some of the challenges faced by managers of intensive pasture systems, but it is unclear how to design the most suitable plant mixtures. Using innovative design theory, we identify two conceptual shifts that foster potentially beneficial design approaches. Firstly, reframing the goal of mixture design to supporting ecological integrity, rather than delivering lists of desired outcomes, leads to flexible design approaches that support context-specific solutions that should operate within identifiable ecological limits. Secondly, embracing, rather than minimising uncertainty in performance leads to adaptive approaches that could enhance current and future benefits of diversifying pasture. These two fundamental shifts could therefore accelerate the successful redesign of intensive pastures.

Resource-use efficiency drives overyielding via enhanced complementarity.

Overyielding, the primary metric for assessing biodiversity effects on ecosystem functions, is often partitioned into "complementarity" and "selection" components, but this reveals nothing about the role of increased resource use, resource-use efficiency, or trait plasticity. We obtained multiple overyielding values by comparing productivity in a five-species mixture to expected values from its component monocultures at a) six levels of nitrogen addition (spanning 0-500 kg N ha-1 year-1) and b) across four seasons. We also measured light, water, and nitrogen use, resource-use efficiency, and three functional traits-leaf nitrogen content, specific leaf area, and leaf area ratio-n mixtures and monocultures. We found strong evidence for non-transgressive overyielding. This was strongest in spring, with mixture productivity exceeding expected values by 20 kg dry matter ha-1 day-1. Peak overyielding was driven by enhanced complementarity, with the two non-N2-fixing forb species far exceeding expected productivity in mixtures. Peak overyielding also coincided with higher water use in the mixture than for any monoculture, and enhanced mixture-resource-use efficiency. There was only weak evidence that trait plasticity influenced overyielding or resource use. Our findings suggest that when complementarity drives overyielding in grassland mixtures, and this is made possible both by increased water use and enhanced efficiency in water, nitrogen, and light use. Our results also suggest that mixtures offer a viable compromise between productivity, resource-use efficiency, and reduced environmental impacts (i.e., nitrate leaching) from intensive agriculture.

Irrigation increases forage production of newly established lucerne but enhances net ecosystem carbon losses.

In New Zealand, dairy farming faces increasing scrutiny for its environmental impacts, including those on soil carbon (C) stocks; hence, alternative management practices are required. One such practice is usage of deep-rooting forage, such as lucerne (Medicago sativa L.). We measured the C and water exchange of two neighbouring lucerne fields on stony, well-drained soil for 3 years, following conversion from grassland. One field received irrigation and effluent; the other received neither. Net CO2 exchange and evaporation were measured by eddy covariance, drainage and leaching with lysimeters, and water inputs with rain gauges. Biomass removal from harvesting and grazing was recorded by direct sampling. In the conversion year, irrigated lucerne was C-neutral despite two harvests and losses from the conversion process. In the 2nd and 3rd years combined, the biomass-C removal exceeded net CO2 uptake, causing net losses of 450 g C m-2 and 210 g C m-2 for irrigated and non-irrigated lucerne, respectively. Leaching losses accounted for 1 to 9 % of annual net C uptake from the atmosphere. The ratio of ecosystem respiration to gross photosynthetic productivity (GPP) increased from <0.7 in spring to ≈ 1 in autumn. Consequently, the net C balance for both lucerne crops showed gains in the first two growth periods of each year and losses in the subsequent two to four growth periods. Irrigation made no difference to the photosynthetic water-use efficiency at field scale (GPP/evaporation), but enhanced production water-use efficiency (biomass/water input). Irrigation increased both the absolute amount of drainage and the fraction of water inputs lost by drainage. In one year, significant summer drainage occurred for the irrigated lucerne. To prevent that, soil-water content should be kept well below field capacity but above the crop's water-stress level. Such practice would likely also help retain soil carbon.

Combining field experiments and predictive models to assess potential for increased plant diversity to climate-proof intensive agriculture.

Agricultural production systems face increasing threats from more frequent and extreme weather fluctuations associated with global climate change. While there is mounting evidence that increased plant community diversity can reduce the variability of ecosystem functions (such as primary productivity) in the face of environmental fluctuation, there has been little work testing whether this is true for intensively managed agricultural systems. Using statistical modeling techniques to fit environment-productivity relationships offers an efficient means of leveraging hard-won experimental data to compare the potential variability of different mixtures across a wide range of environmental contexts. We used data from two multiyear field experiments to fit climate-soil-productivity models for two pasture mixtures under intensive grazing-one composed of two drought-sensitive species (standard), and an eight-species mixture including several drought-resistant species (complex). We then used these models to undertake a scoping study estimating the mean and coefficient of variation (CV) of annual productivity for long-term climate data covering all New Zealand on soils with low, medium, or high water-holding capacity. Our results suggest that the complex mixture is likely to have consistently lower CV in productivity, irrespective of soil type or climate regime. Predicted differences in mean annual productivity between mixtures were strongly influenced by soil type and were closely linked to mean annual soil water availability across all soil types. Differences in the CV of productivity were only strongly related to interannual variance in water availability for the lowest water-holding capacity soil. Our results show that there is considerable scope for mixtures including drought-tolerant species to enhance certainty in intensive pastoral systems. This provides justification for investing resources in a large-scale distributed experiment involving many sites under different environmental contexts to confirm these findings.

The trade-offs between milk production and soil organic carbon storage in dairy systems under different management and environmental factors.

A possible agricultural climate change mitigation option is to increase the amount of soil organic carbon (SOC). Conversely, some factors might lead to inadvertent losses of SOC. Here, we explore the effect of various management options and environmental changes on SOC storage and milk production of dairy pastures in New Zealand. We used CenW 4.1, a process-based ecophysiological model, to run a range of scenarios to assess the effects of changes in management options, plant properties and environmental factors on SOC and milk production. We tested the model by using 2years of observations of the exchanges of water and CO2 measured with an eddy covariance system on a dairy farm in New Zealand's Waikato region. We obtained excellent agreement between the model and observations, especially for evapotranspiration and net photosynthesis. For the scenario analysis, we found that SOC could be increased through supplying supplemental feed, increasing fertiliser application, or increasing water availability through irrigation on very dry sites, but SOC decreased again for larger increases in water availability. Soil warming strongly reduced SOC. For other changes in key properties, such as changes in soil water-holding capacity and plant root:shoot ratios, SOC changes were often negatively correlated with changes in milk production. The work showed that changes in SOC were determined by the complex interplay between (1) changes in net primary production; (2) the carbon fraction taken off-site through grazing; (3) carbon allocation within the system between labile and stabilised SOC; and (4) changes in SOC decomposition rates. There is a particularly important trade-off between carbon either being removed by grazing or remaining on site and available for SOC formation. Changes in SOC cannot be fully understood unless all four factors are considered together in an overall assessment.

Irrigating grazed pasture decreases soil carbon and nitrogen stocks.

The sustainability of using irrigation to produce food depends not only on the availability of sufficient water, but also on the soil's 'response' to irrigation. Stocks of carbon (C) and nitrogen (N) are key components of soil organic matter (SOM), which is important for sustainable agricultural production. While there is some information about the effects of irrigation on soil C stocks in cropping systems, there is a paucity of such studies in pastoral food production systems. For this study, we sampled soils from 34 paired, irrigated and unirrigated pasture sites across New Zealand (NZ) and analysed these for total C and N. On average, irrigated pastures had significantly (P < 0.05) less soil carbon (C) and nitrogen (N) than adjacent unirrigated pastures, with differences of 6.99 t C ha-1 and 0.58 t N ha-1 in the uppermost 0.3 m. Differences in C and N tended to occur throughout the soil profile, so the cumulative differences increased with depth, and the proportion of the soil C lost from deeper horizons was large. There were no relationships between differences in soil C and N stocks and the length of time under irrigation. This study suggests SOM will decrease when pastures under a temperate climate are irrigated. On this basis, increasing the area of temperate pasture land under irrigation would result in more CO2 in the atmosphere and may directly and indirectly increase N leaching to groundwater. Given the large and increasing area of land being irrigated both in NZ and on a global scale, there is an urgent need to determine whether the results found in this study are also applicable in other regions and under different land management systems (e.g. arable).

Leaf economics spectrum-productivity relationships in intensively grazed pastures depend on dominant species identity.

Plant functional traits are thought to drive variation in primary productivity. However, there is a lack of work examining how dominant species identity affects trait-productivity relationships. The productivity of 12 pasture mixtures was determined in a 3-year field experiment. The mixtures were based on either the winter-active ryegrass (Lolium perenne) or winter-dormant tall fescue (Festuca arundinacea). Different mixtures were obtained by adding forb, legume, and grass species that differ in key leaf economics spectrum (LES) traits to the basic two-species dominant grass-white clover (Trifolium repens) mixtures. We tested for correlations between community-weighted mean (CWM) trait values, functional diversity, and productivity across all plots and within those based on either ryegrass or tall fescue. The winter-dormant forb species (chicory and plantain) had leaf traits consistent with high relative growth rates both per unit leaf area (high leaf thickness) and per unit leaf dry weight (low leaf dry matter content). Together, the two forb species achieved reasonable abundance when grown with either base grass (means of 36% and 53% of total biomass, respectively, with ryegrass tall fescue), but they competed much more strongly with tall fescue than with ryegrass. Consequently, they had a net negative impact on productivity when grown with tall fescue, and a net positive effect when grown with ryegrass. Strongly significant relationships between productivity and CWM values for LES traits were observed across ryegrass-based mixtures, but not across tall fescue-based mixtures. Functional diversity did not have a significant positive effect on productivity for any of the traits. The results show dominant species identity can strongly modify trait-productivity relationships in intensively grazed pastures. This was due to differences in the intensity of competition between dominant species and additional species, suggesting that resource-use complementarity is a necessary prerequisite for trait-productivity relationships.

Modelling carbon and water exchange of a grazed pasture in New Zealand constrained by eddy covariance measurements.

We used two years of eddy covariance (EC) measurements collected over an intensively grazed dairy pasture to better understand the key drivers of changes in soil organic carbon stocks. Analysing grazing systems with EC measurements poses significant challenges as the respiration from grazing animals can result in large short-term CO2 fluxes. As paddocks are grazed only periodically, EC observations derive from a mosaic of paddocks with very different exchange rates. This violates the assumptions implicit in the use of EC methodology. To test whether these challenges could be overcome, and to develop a tool for wider scenario testing, we compared EC measurements with simulation runs with the detailed ecosystem model CenW 4.1. Simulations were run separately for 26 paddocks around the EC tower and coupled to a footprint analysis to estimate net fluxes at the EC tower. Overall, we obtained good agreement between modelled and measured fluxes, especially for the comparison of evapotranspiration rates, with model efficiency of 0.96 for weekly averaged values of the validation data. For net ecosystem productivity (NEP) comparisons, observations were omitted when cattle grazed the paddocks immediately around the tower. With those points omitted, model efficiencies for weekly averaged values of the validation data were 0.78, 0.67 and 0.54 for daytime, night-time and 24-hour NEP, respectively. While not included for model parameterisation, simulated gross primary production also agreed closely with values inferred from eddy covariance measurements (model efficiency of 0.84 for weekly averages). The study confirmed that CenW simulations could adequately model carbon and water exchange in grazed pastures. It highlighted the critical role of animal respiration for net CO2 fluxes, and showed that EC studies of grazed pastures need to consider the best approach of accounting for this important flux to avoid unbalanced accounting.