BD5208 Wide Scale Enhancement of Biodiversity (WEB) Final Report on Phase 2, and Overview of Whole Project Executive Summary

BD5208 Wide Scale Enhancement of Biodiversity (WEB) Final report on phase 2, and overview of whole project Executive summary

Core objective

The WEB project aimed to inform the development of new or existing Entry Level (ELS) and Higher Level Stewardship scheme (HLS) options that create grassland of modest biodiversity value, and deliver environmental ecosystem services, on large areas of land with little or no potential for creation or restoration of BAP Priority Habitat grassland.

Specific objectives

 Quantify the success of establishing a limited number of plant species into seedbeds (ELS/HLS creation option) and existing grassland (currently HLS restoration option) to provide pollen, nectar, seed, and/or spatial and structural heterogeneity.  Quantify the effects of grassland creation and sward restoration on faunal diversity/abundance, forage production and quality, soil properties and nutrient losses.  Develop grazing and cutting management practices to enhance biodiversity, minimise pollution and benefit agronomic performance.  Liaise with Natural England to produce specifications for new or modified ES options, and detailed guidance for their successful management.

Overview of experiment: The vast majority of lowland grasslands in the UK have been agriculturally improved, receiving inputs of inorganic fertiliser, reseeding, improved drainage and are managed with intensive cutting and grazing regimes. While this has increased livestock productivity it has led to grasslands that are species-poor in both native plants and invertebrates. To rectify this simple Entry Level Stewardship scheme options have been developed that reduce fertiliser inputs; this includes the EK2 and EK3 options. While permanent grasslands receiving low fertiliser inputs account for the largest area of lowland managed under the agri-environment schemes they currently provide only minimal benefits for biodiversity or ecosystem services. In addition, such low input grasslands have low value for livestock production. Developing new multi-functional seed mixtures and understanding how they can be managed to enhance biodiversity, ecosystem service delivery,

1 soil structure and nutrient value while simultaneously supporting a commercially attractive level of livestock production is a missing link in the toolbox of Entry Level Stewardship grassland options. This Wide Scale Enhancement of Biodiversity project aims to address these issues.

In 2008 a multi-factorial experiment was established to develop simple, low cost (seed cost ≤ £230 ha-1) options that both enhance biodiversity and ecosystem services, including crop pollination and soil structure. We established three seed mixtures differing in the key groups of plants they contained (‘grass’ only (G), ‘grass & legume’ (GL), and ‘grass, legume & forb’ (GLF)), and investigated management practices involved in their establishment and long term maintenance. These included seed bed preparation (‘minimal tillage cultivation’ vs ‘conventional deep ploughing’), the type of sward management (cut vs grazed) and intensity with which that management is applied (typical continuous management vs a summer rested period). The study was undertaken on two experimental grasslands, one in Devon (North Wyke) and one in Berkshire (Jealott’s Hill). These grasslands had previously been agriculturally improved, having been both reseeded as well as receiving regular application of inorganic fertiliser to maximise livestock productivity. North Wyke was last ploughed and reseeded with perennial ryegrass and white clover in 1998. Jeallot’s Hill was previously under arable cultivation in 2004. Although both were floristically species poor (less than 3 species m-2) underlying soil conditions contrasted significantly, with one site on poorly structured compacted London clay (Jealott’s Hill) and the other on brownish clay loams sitting over impermeable clay (North Wyke). Direct monitoring of these experimental sites occurred over four years from 2009 to 2012 following their sowing in 2008.

In this executive summary we highlight our principal findings in terms of how our experimental treatments influenced: 1) The establishment of ecological generalist legume and non-legume forb species into high fertility grassland following conventional or minimal cultivation; 2) sward and livestock productivity; 3) persistence of sown plants over the five years once established into the grasslands; 4) soil parameters, including nutrient loading and soil compaction; 5) water leachate quality as an indicator of diffuse pollution risks; and 6) food web enhancement focusing on insect pollinators and the provision of feeding resources for farmland birds. Summary tables are given after the text.

The establishment of generalist legume and non-legume forbs

The utility of any seed mixture in delivering increased sward productivity, biodiversity or promoting aspects of soil structure and nutrient retention is ultimately dependent on its capacity to establish effectively into nutrient enriched competitive ex-improved grassland swards. This question was addressed in Phase 1 of the study (2008-2010). At North Wyke ploughing and conventional seed bed preparation was the most successful cultivation practice. At Jealott’s Hill the benefits of ploughing over shallow cultivation were equivocal. The results of this study indicate that for shallow cultivation to be successful in allowing high establishment of sown legume and non-legume forb species severe reduction in competition by the existing/original sward constituents is required. At Jealott’s Hill the shallow cultivation technique completely disturbed the existing sward and created more than 80 percent bare ground, whereas at North Wyke the shallow cultivation technique disturbed 40 - 50 percent of the existing sward creating 40-50 percent bare ground. It is probable that if herbicide had

2 been used in conjunction with the shallow cultivation at North Wyke competition by the existing sward would have been controlled and shallow cultivation may have resulted in establishment success comparable to the ploughed treatment.

Productivity

Farmer’s uptake and perception of the value of these seed mixtures will depend to a large extent on their capacity to support livestock, as measured by biomass production and forage quality. Understanding the extent to which different seed mixtures are able to compensate for the loss of inorganic fertiliser inputs is therefore valuable when evaluating the utility of these swards as ELS or HLS options. At North Wyke the dry matter yields of cut silage and the animal stocking rate that the established grasslands were able to support were greatest within the unfertilised more diverse seed mixtures, that is the ‘grass & legume’ (GL) and ‘grass, legume & forb’ (GLF). Although stocking rates were not investigated at Jealott’s Hill, these same diverse seed mixes were found to support higher dry matter yields from the cut silage. Peak dry matter yield from the GL and GLF sward were typically in the range of 6 – 8 tonnes ha-1, although over time the loss of agricultural cultivars of legumes from the sward meant that by 2012 this had been reduced to c. 3 tonnes ha-1 at one site (Jealott’s Hill). This increased yield, at least initially, was directly linked to the establishment of nitrogen fixing legumes within the GL and GLF swards. For example at Jealott’s Hill dry matter yield was c. 5-6 tonnes ha-1 higher in the GLF plots relative to swards established using the ‘grass’ only (G) seed mixture. Although sward productivity was generally higher at North Wyke, poorer establishment and persistence of legumes meant that the difference in dry matter yields between the G and the more diverse GL and GLF seed mixtures was smaller, typically between 1-3 tonnes ha- 1. Where initial seed bed cultivation in 2008 was achieved using minimum tillage, as opposed to more conventional deep ploughing approaches, there was also an indication that silage cuts produced greater dry matter yields, although typically by a small amount of no more than 1 tonne ha-1 for the more productive GLF seed mixes.

The nutritional quality of the forage for livestock was also superior where legumes were included as part of the establishing seed mixtures. Sward nutritional value was higher in the GL and GLF plots where legumes were sown, although at North Wyke non-legume forbs further increased forage quality in the GLF seed mixture. Even though this was not seen at Jealott’s Hill there was evidence that the rate of decline in herbage nutritional quality over time could be reduced where non-legume forbs were also part of the sown seed mixture. In general though this improved nutritional value reflects the key role played by nitrogen fixing legumes, and was most apparent in the higher nitrogen concentrations of forage in the GL and GLF silage. At least at North Wyke, deeper-rooting forb species may have been able to access micronutrients within the soil matrix leading to the increased mineral content for the GLF seed mixtures at this site. As agricultural cultivars of legumes did not persist throughout the entire five year period there tended to be a reduction in the nutritional value of forage over time, for example at Jealott’s Hill forage quality in terms of the content of nitrogen, phosphorus and potassium fell over the five years. Shallow minimum tillage cultivation used to create the seed bed in 2008 increased the nitrogen content of the sward at Jealott’s Hill, although this effect was small. At North Wyke there was also evidence that animal excreta within the grazed plots returned nutrients to the soil acting to fertilise plants within these plots to the benefit of forage quality.

Persistence of sown species

The seed mixtures chosen to establish the swards were intended to be of relatively low cost (≤ £230 ha, ie comparable to the costs of establishing arable field margin options) and so were typically composed of commercially available agricultural cultivars of grasses, legumes and ‘wild flowers’. However, many such cultivars persist poorly once established into swards and so their value over the five year period of a typical Entry Level Stewardship agreement is limited. This is particularly true of many of the legumes which often persist for only 2-3 years, such as Red or Alsike Clover. A failure for this sown component to persist over this period limits their value for biodiversity, particularly in the case of insect pollinators that will forage on these flowering plants. Understanding how best to maximise species persistence once sown is crucial.

The cover and species richness of legumes decreased over the four year period, as agricultural cultivars failed to re-establish by self-seeding into the existing swards. At Jealott’s Hill the rate at which sown legume species were lost from the sward was minimised where a conventional cutting regime involving two cuts per year was applied whereas at North Wyke legumes persisted better under the grazing management. There was some evidence at Jealott’s Hill that under grazing management establishment of legumes was highest where they had been sown as part of the more diverse GLF seed mix. However, establishment of legumes under cutting management did not differ between the GL and GLF seed mixes. At North Wyke the establishment of legume species tended to be highest where they had been sown as part of the more diverse GLF seed mix. However, no evidence was found that the inclusion of non-legume forbs in the seed mix aided the persistence of legume species. The cover of non-legume forbs also fell at both sites over the five year period. This collapse was particularly rapid at Jealott’s Hill, where forb cover in cut plots fell from c. 50 % in the establishment year to around 20 % cover in the following year. However, subsequent to this initial rapid decline from 2009 to 2010 the cover of forbs remained fairly constant under cutting management. While cover declined at both sites, there was evidence at Jealott’s Hill of a small increase in forb species richness over the five years, although this effectively equated to the colonisation of only one additional species into most plots. This colonisation is attributed to a general reduction in competition with legumes whose cover and species richness was declining over this time. At North Wyke the persistence of non-legume forbs showed similarities to the patterns seen at Jealott’s Hill where cutting was the superior management option. Over a five year time scale, sown grasses, legumes and non-legume forbs were of greater abundance and richness at North Wyke where the seed bed had been prepared by ploughing. As already indicated the low persistence of sown legumes and forbs may be due to the type of seed used within the experiment. The agronomic cultivars of these plants are likely to have been bred so that productivity is enhanced at the expense of longevity. A wild-type seed mix may possibly have the opposite trade-off. A mixture of agronomic cultivars and wild-type seeds (if available in sufficient quantity) may lead to an initial boost in plant productivity as the cultivars emerge into the sward, and the wild-type plants persisting in the sward to provide resources for prolonged food web enhancement. However, this has clear limitations in terms of the cost linked with procuring wild type seeds and so may be unsuitable for ELS style options.

Soil parameters

ELS grassland options have the potential not only to mitigate against biodiversity loss, but to improve soil quality in terms of its structure, nutrient status and the rates at which these nutrients are lost from the soils in water leachate. Perhaps the most direct impact on the soil was the cultivation method used to create the seed beds in 2008. The maintenance of a wide range of soil nutrients was highest where minimum tillage cultivation approaches were used, disturbing the soil to a depth of only 5 cm before the seed mixes were added. For example in the top 7.5 cm of soil nitrogen was c. 30 % higher at North Wyke and c. 6 % higher at Jealott’s Hill where minimum tillage cultivation was used relative to conventional deep ploughing in which the more nutrient and carbon- rich topsoil is inverted. At both sites the use of minimum tillage had direct benefits to the stocks of soil carbon. This increase in soil carbon was greatest at North Wyke, which saw a c. 50 - 60 % increase. This has significant implications for the sequestration of greenhouse gasses in grasslands is such options had a wide scale uptake as part of ELS grassland schemes. However, this change in soil carbon may be the result of deep ploughing causing a redistribution of soil carbon from the surface to a depth of c. 20cm as the soil was turned over. As a result there may be no net change in carbon stocks. It is also possible that ploughing and the preparation of a seed bed during a hot dry period of the summer may have removed a significant proportion of the soil organic matter, with a consequent lowering of soil nutrient status. The greater levels of soil nutrients under minimum cultivation when compared to deep ploughing reflect the greater retention of soil nutrients following minimal physical disturbance of the soil in preparing a seed bed. There was some evidence of increased soil pH in the ploughed plots, although it is possible again that this is ultimately linked to the inversion of the soil surface burying more acidic top soils.

Soil phosphorus levels tended to decline over time following the suspension of inorganic fertiliser applications at all sites. While rate of this decline was relatively rapid, it is not outside previously reported drops in phosphorus levels following the suspension of inorganic fertiliser inputs into improved grasslands (Van der Salm et al 2009). There was also some indication of management effects on soil nutrients, although these were largely small. For example, at Jealott’s Hill the deposition of faeces from livestock increased soil carbon stocks, while the removal of biomass as silage under cutting regimes meant that on average carbon decreased over the five years. As this equated to only a 5 % decline in soil carbon under cutting this effect was of limited importance. This impact of livestock on soil nutrient status was also seen at North Wyke where higher levels of total phosphorus and lower soil pH were found in grazed plots. While there was an expectation that diversification of the sward and the extensification of management intensity would have increase the fungal: bacterial ratio in the soil, no evidence of this was found. This suggests that four years is not long enough to shift the soil towards a state of self-regulation in nutrient status.

Soil structure, in particular soil compaction, is a major problem in agricultural systems. At North Wyke soil structure was affected by ploughing, with the top 10 cm of the soil having a greater bulk density in the deep ploughed as opposed to those established using minimum cultivation approaches. However, the pressure required for a probe to penetrate the soil to a depth of 45 cm (a second measure of soil compaction) was reduced where deep ploughing was used, although only to the depth of the plough (20-30 cm). This effect of deep ploughing on soil compaction may reflect the removal of organic matter from the surface layer during the actual ploughing of the plots. The plough when inverting the soil either buried organic matter in the deeper layers of the soil profile or alleviated compaction at these depths. The soil bulk density was also greater where plots were cut rather than grazed during the summer months, presumably due to the machinery trafficking.

Jealott’s Hill was established on far more compacted soils composed of London clay, and while the same cultivation methods were applied we were unable to identify any effect of seed bed preparation on the bulk density of these soils in either the 0-10 or 10-20 cm strata.

The use of plant species with varying rooting characteristics, including some with tap roots, as a mitigation measure to ameliorate soil compaction is one potential benefit of sowing more diverse seed mixtures. At North Wyke the surface (0-10 cm) of the soil had a higher bulk density for the G and GLF seed mixture than was identified in the GL plots. This pattern is counter to the suggestion that deep rooted plants, typical of the forbs sown into the GLF plots, were reducing soil compaction by breaking up the soil. In part, this may be a product of the generally good establishment of red clovers which are moderately deep rooted (Weaver 1926), at this site, particularly relative to many of the more deep rooting non-legume forbs of the GLF seed mix. However, at Jealott’s Hill a reduction in the pressure required to penetrate the soil was found where more diverse seed mixtures were sown (GL and GLF), suggesting that they acted to reduce soil compaction to some extent.

Water leachate

How effective soils are at retaining nutrients has implications not only for local soil fertility, but will also impact on issues relating to diffuse water pollution. At North Wyke, water samples taken from plots where the seed bed was established using minimum tillage cultivation methods were found to have increased levels of total phosphorus in water leachate compared to the deep ploughed plots. However, this may be an artefact resulting from the generally lower levels of soil nutrients found in the surface layer of the deep ploughed plots. However, this explanation poses the question of why more total oxidised nitrogen did not leach from the minimum cultivation plots given the higher amounts of total nitrogen available in the soil. Over a five year time scale deep rooting plants did not reduce total phosphorus leached at North Wyke as higher concentrations were seen in the more diverse seed mixtures. Introducing a proportion of wild-type species into the seed mix may increase the persistence of forbs thus forming a more substantial root network which may aid with reducing nutrient loses.

Food web enhancement

Increasing the diversity and cover of plants within the swards using sown seed mixtures will provide key food resources for higher trophic levels that would otherwise find agriculturally improved grasslands hostile habitats. At both sites the availability of pollen and nectar resources utilised by insect pollinators was strongly linked to the establishment and persistence of sown legumes and forbs. Legume flower counts were as high as c. 80 m-2, although this fell rapidly over the five year period so that by 2012 typically less than 10 m-2 were found. This contrasted with the density of non-legume forb flowers which increased over the same time period from c. 5 to 20 m-2. At Jealott’s Hill cutting management, particularly where only a single yearly sward cut occurred, supported the greatest densities of both forb and legume flowers. This pattern was repeated at North Wyke for the legumes, but more forbs were recorded at this site within the grazed plots. This may reflect the grazing preferences of the cattle used, with some forb plant species (e.g. Chicory and Lesser Knapweed) being avoided once mature. At both sites the use of deep ploughing to create the seed bed in 2008 resulted in the highest flower densities for both legumes and forb flowers.

The occurrence of insect pollinators (bees, butterflies and hoverflies) closely tracked the availability of their foraging flower resource. This meant that over the five year period there was a general pattern of declining abundance and species richness of pollinators as the flowers they fed on were lost from both sites. Although the total abundances of pollinators in the GL and GLF seed mixes when managed by a single silage cut were at a similarly high level in 2009, by the final year their abundance had fallen by c. 90 % in the GL plots but by only c. 30 % in the GLF plots. This robustness of the pollinator community in the GLF seed mixture was linked to the density of forb flowers which increased over time at this site and so compensated for the loss of legume flowers over the same period. At North Wyke the initial establishment of forbs was poorer, and as a result their flower densities declined over the five years and so were unable to compensate for a similar loss in legume flower density.

For both North Wyke and Jealott’s Hill the method of cultivation used to establish the seed beds influenced the amount of seed heads available for birds to feed on over the autumn and winter period. At North Wyke grasses and legumes were more abundant in minimally cultivated plots whereas forbs were more abundant in ploughed plots. For Jealott’s Hill seed head densities of both legume and forb seeds were highest where deep ploughing was used to establish the plots. There was also some evidence that seed heads may be more abundant in the grazed as opposed to the cut plots, at least at the North Wyke site. However, in the case of the legumes seed head density over the winter was dictated by the timing of the management. A cessation of summer grazing or the absence of a late season sward cut was crucial to supporting high abundances of seed head over the winter. Management that continued throughout the summer acted to deplete this resource for winter feeding birds, whether the swards were managed by cutting or grazing. This importance of a management rest period is linked to providing the plants with a sufficient period of time to reach full phenological development.

For most farmland birds insects are a crucial component of their diets, particularly when rearing chicks. At both North Wyke and Jealott’s Hill the biomass of beetles, representing a key food resource for insectivorous birds, was significantly higher in swards established using the more diverse GL and GLF seed mixes. This was linked to the strong feeding associations of many plant species with the sown wildflower components, in particular legumes. For the establishment year at Jealott’s Hill, the biomass of beetles supported in the unfertilised G plots was around only 5 – 10 % of that in the floristically diverse GL and GLF plots. In general, rested management (a single sward cut or no summer grazing) supported the highest biomass of beetles, and cutting management was superior to grazing. This importance of rested management reflects the need for a period of no disturbance within which populations of the beetles were able to increase. However, the biomass of beetles supported in the plots decreased in response to the loss of legumes and forb cover, so that as much as c. 75% of the beetle biomass was lost over the five years. Policy notes

A number of recommendations can be made from the experimental findings. For the analyses that support these recommendations see sections Part 1 and 2 below.

Seed bed preparation: The method used for seed bed preparation has direct benefits in terms of above ground biodiversity and below ground soil nutrient retention.

 Successful establishment of forbs and legumes was superior following deep cultivation combined with a herbicide application compared with surface cultivation without herbicide. Reduced levels of competition as the existing sward was buried combined with the removal of fast growing weed species had direct benefits for the establishment of the sown floral component of seed mixtures. The good establishment of forbs and legumes has benefits for those species that feed upon them, directly or indirectly.  Minimum tillage combined with herbicide application may reduce competition and promote the establishment of the sown seed mixtures. Although not tested, this compromise approach could have the benefits of ensuring good establishment and maximising soil nutrient status. Alternatively minimum tillage approaches could be exaggerated so that more of the soil surface is disturbed, that is to say greater than the 40 % cover disturbance aimed for in this study. By not disrupting the sub-soil (>5 cm depth) 70- 90 % soil disturbance may still allow high levels of forb and legume establishment while protecting the soil nutrient status.

Seed mixture: Although more expensive (grass only = £100 ha-1 vs. £140-230 ha-1) the addition of legumes and non-legume forbs to seed mixes represent the most important finding of this study. This has direct benefits in several areas of biodiversity, ecosystem service provision and livestock production. These are:

 Herbage biomass, nutrient quality and livestock production: Swards sown with both legumes and non-legume forbs have increased dry matter yield, forage quality and animal performance compared to non-fertilized grass-only swards.  More diverse seed mixtures may reduce some aspects of soil compaction: Sowing both legumes and forbs was shown to reduce the force needed to penetrate soils, although this effect may be more pronounced on highly compacted soils.  Modest but wide scale enhancement of floristic diversity using simple seed mixtures was shown to dramatically increase the resource base of flowering plants as well as their utilisation by insect pollinators. This would be likely to lead to positive population growth for a suite of insect pollinators in agricultural land, although other caveats like the availability of nesting sites for bees would be an issue. Particularly in the context of mixed agricultural systems this may benefit the delivery of pollination services that contribute to increased yields of mass flowering crops, soft and top fruit.  Including both legumes and non-legume forbs within seed mixtures increases the persistence of flowering resources for insect pollinators. The rapid loss of agricultural cultivars of legumes from the sward can be compensated by a modest increase in forb flower density over the typical five year agreements associated with ELS.  The cheaper cultivars of red clover and other legumes, which are most likely to be sown, will only persist for 1-2 years after the establishment year. This means that, in order to sustain the resource for invertebrates and the other benefits, they will need to be resown. An alternative would be to sow more persistent but more expensive wild types.

Sward management: Although the establishing seed mixture is fundamental in ensuring that ELS options linked with improved grasslands deliver on a suite of biodiversity and ecosystem service

8 goals, subsequent management will have implications for the persistence and scale of delivery of these factors.

 Cutting management regimes tend to promote persistence of legume and forb populations, and so lead to higher numbers of pollinators than occurs in grazed areas. However, a mixed management strategy has benefits, as the cutting events totally removed all flower resources for a short period of time. The more gradual impact of grazing, particularly when suspended over the summer, allowed for a more continuous foraging resource for insect pollinators albeit at a lower overall density of flowers.  Rested management practices support more feeding resource for insectivorous farmland birds over the summer. A rest period in the mid-summer leads to greater availability of beetle biomass that can be fed on by birds. It is worth noting that access to this biomass by birds is likely to be easiest in grazed plots, as typically the height of the cut plots was more uniform.  Seed resources for overwintering farmland birds are superior where grazing or cutting is suspended in the mid and latter part of the season. A summer suspension of grazing or the avoidance of a late season sward cut is vital to allow a rest period where seed heads can develop and persist as a resource for winter feeding birds.

Future work

Potential directions for future work can be considered to fall into two spatial sales which: 1) identify further refining of management operations at field scales to promote the successful delivery of resources and; 2) consider how the implementation of these approaches at landscape scales will best deliver biodiversity and ecosystem service goals in a cost effective manner.

Field scale studies:

 Examine the potential for including with a mix of agronomic cultivars, wild-type seeds of sown species to provide a win-win with greater productivity and persistence for greater food web enhancement.  Investigate the potential for integrating WEB type swards into dairy systems  Examine the potential of using WEB type swards with arable rotations to enhance soil conservation and expand whole farm Ecosystem Service provision.  Optimise the potential values of using different forb species to achieve nutritional benefits for livestock and other Ecosystem Services including reduction of methane emissions and hence mitigation of climate change.  Investigate what impact WEB type grasslands have on below-ground biodiversity and function.  Investigate why the inclusion of forbs may improve establishment and persistence of legumes.

 Develop minimal cultivation practice(s) to achieve high establishment and persistence of WEB type mixtures, which also provides a high level of soil conservation and minimal nutrient loss.  Design a tool-box for land managers to provide cost-benefits of WEB type grasslands: for agronomic and the provision of Ecosystem Services, such as resources for pollinators and over-wintering birds.  Improve understanding of relationship between rooting depth/morphology of plant species and soil structure.

Landscape scale studies:

 One of the key outcomes of this research is the potential use of floristically enhanced grasslands to support and mitigate against declines of insect pollinator populations in agricultural systems. While we have identified the value of using legume and non-legume seed mixes to enhance the resource value of these grasslands, our understanding of the scale at which they need to be implemented within landscapes to support insect pollinators is limited. Further research is required to identify the impacts of scaling issues on the proportion of land and its spatial arrangement on the resource value of these grassland systems for pollinators. The implementation of these seed mixes as part of ELS options via the point allocation system could be used to affect farmer uptake, and so their implementation at landscape scales.  As a related research question the value of insect pollinators as providers of this ecosystem service will be affected by their ability to forage large distances into crops. Research that directly identifies how proximity to these newly created grasslands will promote increased spill over of the pollinators into the crop, and to what extent this translates into increased yields is required.

Current academic outputs

The following peer reviewed articles leading from work undertaken as part of this project are either currently published or in an advanced state of preparation.

Pywell R.F., Woodcock B.A., Tallowin J.R., Mortimer S.R., Bullock J.M. (2012) Restoring species-rich grassland: principles and techniques. Aspects of Applied Biology (115): Restoring diverse grasslands: what can be achieved, where and what will it do for us?

Woodcock, B.A., Bullock, J.M., Nowakowski, M., Orr, R., Tallowin, J.R.B., & Pywell, R.F. (2012) Enhancing floral diversity to increase the robustness of grassland beetle assemblages to environmental change. Conservation Letters, 5:459-469.

Woodcock, B.A., Savage, J., Bullock, J.M., Nowakowski, M., Orr, R., Tallowin, J.R., & Pywell, R. (2013) Enhancing beetle and spider communities in agricultural grasslands: the roles of seed addition and habitat management. Agriculture Ecosystems & Environment, 167: 79-85. Providing foraging resources for pollinators in UK improved grassland systems

Woodcock, B.A., Savage, J., Bullock, J.M., Nowakowski, M., Orr, R., Tallowin, J.R.B. & Pywell, R.F. (Submitted) Providing foraging resources for pollinators in UK improved grassland systems. Biological Conservation.

Orford, K.A., Murray, P.J., Memmott, J. (Submitted) Engineering biodiversity: Manipulation of plant species richness and grassland management to increase pollinator community functioning.

Summary of general findings at North Wyke (read in conjunction with the text). Cultivation Seed mix Management Timing MC P G GL GLF Cut Graze Typical Rested DM Yield o o o + + N/A + - Forage quality + - - + + - + N/A

Productivity Animal performance N/A - + + N/A N/A Sown grass + + + + o + - o o Sown legume - N/A + - + -

Plant Plant + + +

persistence Sown forb - + N/A N/A + + - + - Total P + - - + o - + N/A Olsen P + - - + - o o N/A Total C + - - +/o - o o N/A Total N + - - +/o - o o N/A pH - + o o o + - N/A Bulk density 0-10cm + - - + - - + N/A

Soil chemistry andSoil structure chemistry Bulk density 10-20cm - + + + + - + N/A TON o - o o - - + N/A

Water TP + + + + + + - N/A Pollinator abundance + + o + + + - + - Legume inflorescence (no.) + + N/A + + + - + - Forb inflorescence (no.) + + N/A N/A + - + o o Seed-head availability G + o + + o - + - + " " L + + N/A + + - + - + " " F - + N/A N/A - + Food web enhancement Food web + - + Beetle biomass + + - + + + - - + Note: + denotes a positive effect, - denotes a negative effect, o denotes a neutral effect, and size of the + or - illustrates the strength of the effect.

Summary of findings at Jealott’s Hill Cultivation Seed mix Management Timing Shallow Deep G GL GLF Cut Graze Typical Rested DM yield + - - + + N/A N/A + - Productivity Forage quality + - - + + N/A NA + - Animal Performance† N/A N/A N/A N/A N/A N/A N/A N/A N/A Sown grass o o + o o o o + - Plant Sown legume o o N/A o o - + persistence + + Sown forb - + N/A N/A + o o - + Total Phosphorus + - o o o o o N/A N/A

Olsens Phosphorus o o o o o o o N/A N/A

Total Carbon + - o o o X X N/A N/A Total Nitrogen + - o o o o o N/A N/A

structure pH o o o o o o o N/A N/A

Bulk density (0-10cm) o o o o o o o N/A N/A Soil chemistry and and chemistry Soil Bulk density (10-20 cm) o o o o o o o N/A N/A PLFA Fungal: bacterial ratio o o o o o o o N/A N/A Water ‡ Total Oxidized Nitrogen / Phos. N/A N/A N/A N/A N/A N/A N/A N/A N/A Pollinator abundance o o - + + + - - + Legume inflorescence (density) o o N/A + + + - - + Forb inflorescence (density) o o N/A N/A + + - - + Seed head availability G o o + - - + - - + “ “ “ L - + N/A + + X X - + “ “ “ F - + N/A N/A + - + - +

Food web enhancement web Food Beetle biomass + - - + + o o - +

Note: Symbols size denotes the strength of positive (+), negative (-) and neutral effects (o); X = effects that are confounded by interaction terms and so do not have simple responses to a treatment; G = Grasses; L = legumes; F = non-legume forbs; N/A = comparisons either not applicable or not made for this site. Note (†) that animal performance measures were only undertaken at North Wyke and (‡) water measurements proved impossible due to the compact clay soils of Jealott’s Hill.

Methods

As already described within the executive summary, the overall aim of the WEB project was to inform the development of new or existing Entry Level (ELS) / Higher Level Stewardship schemes (HLS) that would create grassland of modest biodiversity value on large areas of land unsuitable for BAP priority habitats. This aim was split into four specific objectives that dealt with the outcomes of the research and how we would interface into the regulatory bodies. These specific objectives which were: 1. Quantify the success of establishing a limited number of plant species into seedbeds (ELS/HLS creation option) and existing grassland (currently HLS restoration option) to provide pollen, nectar, seed, and/or spatial and structural heterogeneity.

2. Quantify the effects of grassland creation and sward restoration on faunal diversity/abundance, forage production and quality, soil properties and nutrient losses.

3. Develop grazing and cutting management practices to enhance biodiversity, minimise pollution and benefit agronomic performance.

4. Liaise with Natural England to produce specifications for new or modified ES options, and detailed guidance for their successful management.

To address these objectives seven key hypotheses were developed to underpin scientific goals and dictate the experimental design, analysis and interpretation of the data. These Hypotheses were:

H1: Ecological generalist legume and non-legume forb species can be readily established in high fertility grassland following conventional or minimal cultivation. Note that H1 was addressed in the phase 1 report for the Wide Scale Enhancement of Biodiversity (BD1466) project and so discussion of its results will not be repeated in the Phase II report.

H2: Conventional seed bed preparation will result in major nutrient leaching loss whereas minimal cultivation will result in negligible nutrient loss during the establishment and subsequent management of the legume or legume plus forb rich swards.

H3: Sown ecological generalist legume and non-legume forb species can regenerate and persist for at least three years under standard/typical grazing or cutting regimes or summer rested regimes.

H4: Establishment of non-legume forbs will incrementally increase the abundance and diversity of pollen and nectar feeding invertebrates compared with grass-dominated or legume rich swards.

H5: Beneficial effects of legume rich or legume and forb rich swards on pollen and nectar feeding invertebrates will be increased when the swards are rested from grazing or cutting in the mid- to late-summer period.

H6: Establishment of deep rooted species will enhance soil structure and reduce nutrient loss via leaching under either grazing or cutting management systems.

H7: Legume or legume plus forb rich swards will increase grassland productivity and forage quality compared with very low input grass dominated swards.

The methodologies and approached used to test these seven hypotheses are outlined below.

Experimental sites and design

Field-scale, multi-factorial experiments were established in the summer of 2008 at two sites: 1) North Wyke, Devon (SX645990) on brownish clay loams sitting over impermeable clay soil of the Hallsworth series in an area of high winter rainfall. This site had a history of pasture management since the early 1980s, was last ploughed and reseeded with a Lolium perenne and Trifolium repens seed mix in 1998 and had a soil P index of 2. Detailed soil descriptions and infiltration rates of the unsown control plots are reported in Appendix 3 and 4. 2) The second site was at Jealott’s Hill at the Syngenta farm in Berkshire (SU883722). This site is situated on highly compacted London clay soils covered unevenly with Plateau Gravel that is occasionally cemented together with iron oxides. This site was under arable cultivation in 2004, and for at least 10 years prior to this. Both sites were originally dominated by Lolium perenne and Trifolium repens ley, 10 years old at North Wyke and < 5 years old at Jealott’s Hill. Results for the 2009 and 2010 field seasons have been published (BD1466).

At both sites a nested randomised block design with four replicates was used to investigate the effects of the following factors on both biodiversity and the delivery of ecosystem services (ES):

1) seed mixture composition (main treatment); 2) type of management (sub-treatment); 3) timing of management (sub-sub-treatment); 4) seed bed preparation (sub-sub-sub-treatment).

The experiment consisted of a total of 18 treatments at North Wyke (Fig.1) and 26 treatments at Jealotts Hill (Fig.2). The latter site represented a balanced experimental design where all factorial combinations of the four experimental treatments were implemented. At the lowest level of the spilt-split-split plot hierarchy plot size under cutting management was typically 0.06ha, while under grazing management were 0.11ha reflecting the need to support livestock in fixed plots at North Wyke.

Grass only Grass + legume Grass + legume + forb

Graze T T T R T R T T R

E† P MC MC MC MC P MC MC

Cut T T T R T R T T R

E† P MC MC MC MC P MC MC

T=Typical management R= Rested management †E= Existing sward P=Plough MC=Minimal Cultivation

Fig. 1. Experimental design/layout of one of the four replicate blocks at North Wyke

Treatment number (1-26) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26

Existing grass Existing GRASS GRASS+LEGUME GRASS+LEGUME+FORB Establishing seed mix (WP)

Graze Cut Management (SP)

Timing of Management (SSP) Typical Rest Shallow Deep Cultivation (SSSP)

WP = Whole plot, SP = Split plot, SSP = split-split plot and SSP = split-split-split plot. The two existing grassland plots were managed using either grazing or cutting management with the timing of operations typical of commercial farming practice. Note that at North Wyke the sub-set of treatments comprising GL established by deep cultivation was not present also there were no ploughed rested treatments, there was, nevertheless, a balanced experimental design in all analyses (see details in statistical section below).

Fig. 2. The balanced split-split-split plot design of Jealott’s Hill

Seed mixtures: At the whole plot level of the experimental design three seed mixtures were sown (Table 1). These were: 1) grass only mix (G) comprising 5 high agronomic performance grass species sown at 30 kg ha-1 at a seed cost of ca. £100 ha-1. 2) grass + legume mix (GL) comprising the 5 grasses used in G and 7 agricultural legumes sown at 34 kg ha-1 at a seed cost of ca. £145 ha-1). 3) grass +

16 legume + non-legume forb mix (GLF) comprising the 5 grass species of G, the 7 legume species of L and 6 non-leguminous forbs, selected on the basis of their multifunctional value (from Mortimer et al., 2006), sown at 33.5 kg ha-1 at a total seed cost of ca. £230 ha-1. The 5 grass species used in the G mixture were: perennial ryegrass (Lolium perenne), cocksfoot (Dactylis glomerata), timothy (Phleum pratense), meadow fescue (Festuca pratensis), and meadow foxtail (Alopecurus pratensis). The legumes used in the GL mixture were: red clover (Trifolium pratense), white clover (Trifolium repens), alsike clover (Trifolium hybridum), birdsfoot trefoil (Lotus corniculatus), sainfoin (Onobrychis viciifolia), ribbed melilot (Melilotus officinalis), and black medick (Medicago lupulina). The 6 non- leguminous forbs used in the GLF mixture were: chicory (Cichorium intybus), salad burnet (Sanguisorba minor), yarrow (Achillea millefolium), lesser knapweed (Centaurea nigra), oxeye daisy (Leucanthemum vulgare), and common sorrel (Rumex acetosa). The G plots were sprayed with a selective herbicide to remove broad-leaved species during the first establishment year.

In addition, there was an unsown ‘existing grassland’ control nested within the grass only main treatment which was considered to be representative of low input semi-improved grassland typical of that under ELS agreement, i.e. EK2 receiving 50kg ha-1 yr-1 N in spring. These plots were dominated by L. perenne and T. repens and were managed either by a typical cutting regime for silage (two silage cuts a year) or by typical grazing by cattle (from April/May to October). The Existing sward plots were not sprayed with a selective herbicide to remove broad leafed species, as the existing sward was intended to be representative of low input semi-improved grassland typical of that under ELS agreement. Although the mean values of these reference communities in the Existing plots are presented in tables within this report for WEB phase 2 (Appendix 1 and 2), they have been excluded from the main analysis for both sites as they were additions to the overall design.

Sowing rates (kg ha-1) used to establishing seed mixtures Grass Grass Grass, only & legume legume & forbs Sown grasses Provenance Alopecurus pratensis Commercial 5.00 3.75 3.00 Dactylis glomerata Commercial (var. Sparta) 6.50 4.88 3.90 Festuca pratensis Commercial (var. Cosmolit) 5.00 3.75 3.00 Lolium perenne Commercial (vars. Aberstar & Orion) 13.00 9.75 7.80 Phleum pratense Commercial (var. Promesse) 0.50 0.38 0.30

Sown Legumes Lotus corniculatus Commercial (var. Giada) 1.00 1.00 Melilotus officinalis Commercial 2.00 2.00 Onobrychis viciifolia Commercial 4.50 4.50 Trifolium dubium Commercial (var. Virgo Pajbjerg ) 1.00 1.00 Trifolium hybridum Commercial (var. Dawn) 1.00 1.00 Trifolium pratense Commercial (vars. Altaswede & Milvus) 1.10 1.10 Trifolium pratense Native (Essex) 0.50 0.50 Trifolium repens Commercial (var. S183) 0.40 0.40

Sown Forbs Achillea millefolium Commercial 0.40 Centaurea nigra Native 0.30 Cichorium intybus Commercial (var. Puna) 1.00 Leucanthemum vulgare Native 0.20 Rumex acetosa Native 0.10 Sanguisorba minor Commercial 2.00 Table 1. Summary composition of sown plant seed mixtures.

Management: The seed mixture main plots were managed either by: 1) moderate cattle grazing at c. three livestock unit ha-1, or 2) silage cutting to a height of 10 cm with subsequent removal of all cut herbage. This represented the first split of the experimental design. Note, that with the exception of 2009 all experimental plots were grazed by sheep in March following standard sward management practices typically employed by farmers.

Timing of management application: Each management sub-treatment was further split into either 1) ‘typical’ or 2) ‘summer rested’ sub-sub-treatments (Table 2a & b). Timing effectively represents a measure of management extensification applied to both the grazing and cutting practices. Typical management represented practice typical of commercial farming. Rested management involved a period in mid-summer with no disturbance either by cutting or grazing to allow forbs to flower. Typical grazing comprised of moderate continuous grazing with cattle from April/May to October. Rested grazing required moderate grazing from April/May - early June, no grazing from early June - August and moderate grazing with cattle from August – October. Typical cutting management was achieved by cutting for silage with a first cut in late May/early June and a second cut in July/August. There was no July/August cut in the rested treatment. The timing of the rested period was intended to allow a window for the phenological development of flower and seed heads in the sown component of the seed mix. Note that some exceptions to these management timings occurred (see below table detailing management timings). In 2009 grazing was initiated relatively late at both sites due to the grass only plots being sprayed with a selective herbicide to remove broad-leaved species. Following spraying no stock could be introduced to these plots for two weeks, hence the herbage became too tall to graze and the plots were cut delaying turnout. There were also issues in sourcing cattle at Jealott’s Hill. Due to drought conditions in 2011 at North Wyke from June instead of G, GL and GLF treatments being continuously stocked, paddocks were rotationally grazed per block. The wet summer of 2012 resulted in a cattle being temporarily removed in mid-August and later first and second silage cuts. At Jealott’s Hill in the establishment year of 2009 three sward cuts were taken due to the unexpectedly high levels of sward growth in all plots. In contrast in 2012 only a single sward cut was taken due to a combination of a very wet year and an overall reduction in sward fertility leading to poor overall sward development. For this reason the difference between rested and typical cut treatments in 2012 is a result of cutting management in previous years. In 2011 drought conditions meant that cattle were removed from all plots in September rather than October.

Cutting Grazing 2009 Typical: 29/5/2009, 10/8/2009 and Typical: 9/6/2009 until 14/10/2009 2/10/2009. Rested: 9/6/2009 until 7/7/2009:

Rested: 29/5/2009 and 2/10/2009. 13/8/2009 until 14/10/2009

2010 Typical: 5/6/2010 and 8/9/2010 Typical: 12/4/2010 until 7/10/2010 Rested: 5/6/2010 Rested: 12/4/2010 until 4/6/2010; 3/8/2010 until 7/10/2010 2011 Typical: 7/6/2011 and 12/9/2011 Typical: 10/4/2011 until 15/9/2011 Rested: 7/6/2011 Rested: 10/4/2011 until 1/6/2011; 29/7/2011 until 15/9/2011 2012 Typical: 28/8/2012 Typical: 27/4/2012 until 27/10/2012 Rested: 28/8/2012 Rested: 27/4/2012 until 16/7/2012; 10/8/2012 until 27/10/2012 Table 2a. Summary of the timing of sward management at Jeallot’s Hill

Cutting Grazing 2009 Typical: 1/6/2009 and 18/8/2009 Typical: 11/6/2009† until 28/9/2009 Rested: 1/6/2009 and 8/9/2009. Rested: 23/6/2009 until 8/9/2009 (grazing excluded) 2010 Typical: 3/6/2010 and 16/8/2010 Typical: 26/4/2010 until 18/10/2010 Rested: 3/6/2010 and 13/9/2010 Rested: 18/6/2010 until 6/9/2010 (grazing excluded) 2011 Typical: 31/5/2011 and 22/8/2011 Typical: 18/4/2011 until 17/10/2011 Rested: 31/5/2011 and 9/9/2011 Rested: 16/6/2010 until 12/9/2010 (grazing excluded) 2012 Typical: 24/7/2012 and 6/9/2012 Typical: 16/5/2012 until 13/8/2012; 10/9/2012 Rested: 24/7/2012 and 8/11/2012 until 5/10/2012 Rested: 15/6/2012 until 10/9/2012 (grazing excluded) † Areas due to be grazed were also cut 1/6/2009 as a consequence of the withdrawal period following spraying to remove white clover from Grass plots.

Table 2b. Summary of the timing of sward management at North Wyke

Cultivation for seed bed preparation in 2008: To assess the effects of initial seed bed cultivation in the autumn of 2008 each sub-sub treatment was divided into either: 1) herbicide application (Glyphosate sprayed at 5 l ha-1 in 100 litres of water ha-1 (360 g a.i. l-1) followed by inversion tillage created using a conventional reversible plough turning soil to a depth of 25-30 cm. The herbicide treatment was used to kill the existing sward while deep ploughing turned over the existing seed bank; 2) a minimum tillage approach where shallow cultivation alone without the use of a herbicide was applied to a depth of c. 5 cm to create a seed bed. At Jealott’s Hill the shallow cultivation technique completely disturbed the existing sward and created more than 80 percent bare ground using a tractor drawn disc harrow, whereas at North Wyke a slot seeder was used which disturbed 40 - 50 percent of the existing sward creating 40-50 percent bare ground. . In both cases, following seed bed preparation, the seed mixtures were broadcast onto the plots and then rolled. A molluscide was applied during the early establishment phase. While the minimum tillage approach has low fuel requirements, uses no herbicides and helps maintains soil carbon stocks, it is relatively ineffective in terms of controlling competitive weed species (Edwards et al 2007; Morrisa et al 2010). This was expected to limit the window of opportunity for small seeded slow-growing wildflowers to establish (Walker et al 2004; Edwards et al 2007).

Measurements

Agronomic measurements:

Grazing stocking rate and forage quality The number of animals and duration over which the swards would support livestock were recorded for each of the separately fenced groups of sub plots on each sown seed mix. This was done to provide an overall measure of: 1) grazing day total on each sward type for the typical treatments and; 2) a measure of the agronomic penalty in terms of lost grazing for the rested treatment. As Jealott’s Hill was operated under an open grazing system (where cattle had access to all grazed plots) this was only undertaken at North Wyke. During the early summer grazing period and the mid- to late-summer period snip samples of herbage were taken from each grazed sub-plot for mineral (N, P, Ca, Mg and Na) and pepsin cellulose digestibility analyses to coincide with the cut herbage assessments at North Wyke. Determination of total nitrogen was by use of the Leco FP 428 nitrogen determinator. Determination of extractable phosphorus in herbage was by colorimetry, and that of sodium, potassium, calcium and magnesium by ICP-AES.

Cutting forage yield and quality To assess the yield of herbage produced during a silage cut swaths left after the tractor cut were weighed and sampled from a known area to provide a measure of dry matter yield. Samples were also taken to assess forage quality (mineral content and digestibility as in the pluck samples). Mineral (N, P, Ca, Mg and Na) and pepsin cellulose digestibility analyses were undertaken for these samples. Determination of total nitrogen was by use of the Leco FP 428 nitrogen determinator. Determination of extractable phosphorus in herbage was by colorimetry, and that of sodium, potassium, calcium and magnesium by ICP-AES.

Biodiversity measurements

Botanical composition Cover of all vascular plant species were estimated yearly in all treatment plots from 2009 to 2012 using the same approach as Phase 1 of the project. In each experimental plot a total of ten 1m × 1m quadrats were placed at points perpendicular to a mid-line transect, with placing along and out from the mid-line chosen at random (Fig. 3). Different random numbers were used for each plot and in each year. This will avoid underlying patterns of soil or vegetation compromising the independence of individual samples. Quadrats were placed alternately on each side of the mid-line, with cover estimations alternating with frequency estimations on each side. Each quadrat was placed in the same manner to avoid bias, e.g. using a compass direction, with quadrats to the west of the mid-line having the south east corner touching the sample point, and quadrats on the east side of the mid- line having the south west corner touching the sample point chosen at random.

Of these ten quadrats, five were (labelled C on Fig. 3) used to estimation percentage cover of vascular plants by vertical projection, as well as the cover of dead vegetation and bare ground. The remaining five quadrats were used to determine the presence or absence of plant species to be combined with data from the other 5 quadrats (labelled F on Fig.3). Note this data presence absence data is not presented in the current report. Botanical surveys were carried out in mid-July to enable rested sub-plots to have re-grown sufficiently to express any differences to those of the typical management treatment.

Fig.3. Representation of botanical sampling of the core area (500 m2) of each plot.

Flower head surveys One of the key aims of the project was to enhance floristic diversity of the swards in terms of legumes and non-legume forbs. In the case of insect pollinators unless the plants are able to reach full phenological development and so flower the resource value of these grasslands would be likely to be minimal. To determine the availability of flower heads for each of the sown species of legumes and non-legume forbs three 0.5 × 0.5 m quadrats were placed at 5, 15, 25 m spacing along two parallel transects each measuring 2 × 30 m and sited through each plot with a separation of c. 6 m, as illustrated below:

In each quadrat all flowering dicot species were identified and counts made of the number of flower units of each species. A flower ‘unit’ consisted of: a head (e.g. Trifolium spp.), a spike (e.g. S. minor) or capitulum (e.g. C. nigra). Each transect was recorded 3 times per year, in mid-Jul, late-Jul and mid- Aug. to coincide with the counts of pollinating insects.

Seed head counts For overwintering farmland birds, seed heads on plants established within the swards represent a potentially important food resource (Holland et al 2006; Mortimer et al 2006). To assess the value of this resource counts were carried out soon after the last grazing and cutting of the season, typically mid- to end- of September. Counts were made along 2 parallel transects in each plot following the method used for pollinator recording (see above). Each transect measured 20 m in length and be were separated by approx. 6 m, as for the flower resources surveying. In each of these transects, three 0.5 × 0.5 m quadrats were placed at the following paced distances along each transect: 5, 10, 15 m, giving a total of 6 quadrats per plot. In each quadrat the number of seed head units (i.e. reproductive structures containing ripe seeds) of each species of grass, legume and non-legume forb were counted. One seed head ‘unit’ is a panicle (grasses), an umbel (e.g. Heracleum sphondylium), a

21 head (e.g. Trifolium spp. or O. viciifolia), spike (e.g. M. officinalis), capitulum (e.g. C. nigra, C. intybus) or individual pod (e.g. L. corniculatus).

Pollinating insects In each experimental plot, two fixed 20 m × 2 m transects were established. Each transect was surveyed for insect pollinators on three separate occasions yearly from 2009 – 2012. Surveys were undertaken between 10.00 - 16.00 hours when temperatures were greater than 13oC in clear conditions (at least 60% of the sky being clear) or greater than 17oC in more cloudy skies (Pollard & Yates, 1993). The three pollinator surveys occurred yearly in mid-May before the first sward cut, late July c.4-6 weeks after the first cut, and early August before the final sward cut. Pollinators were identified at the following taxonomic resolution: 1) All butterflies to species; 2) Honeybee (Apis mellifera); 3) Bumblebees to species Bombus lapidarius, B. terrestris / lucorum, B. pascuorum, B. pratorum, B. hortorum, B. hypnoroum, B. vestalis, B. rupestris and the parasitic sub-genera Psythirus spp.; 4) Total abundance of all solitary bees and hoverflies were made. The plant species on which each insect was first seen foraging was recorded to species. These were used to derive a measure of total pollinator abundance and pollinator species richness for each plot, the latter being composed of the bees and butterflies only. All abundances and species richness are expressed as summed values for all transects for a single year (i.e. based on three transect walks).

Sward active beetles Adult phytophagous beetles were sampled using a Vortis suction sampler (Burkland Ltd, UK) from 2009 to 1012 (see Woodcock et al 2012 and Woodcock et al 2013 for currently published work relating to the work undertaken in the WEB project). Samples were taken on dry days in June and September, each representing 55 × ten second suctions per plot (Brook et al. 2008). All phytophagous beetles (Chrysomelidae, Curculionoidea, Apionidae, Bruchidae, Elateridae and Oedomeridae) were identified to species. Total biomass of each species was also determined to provide an indication of potential energy flow in the form of feeding resources to higher trophic levels, in particular birds (Atkinson et al. 2004; Saint-Germain et al. 2007;). Individual species biomass was determined from body length vs. mass relationships (Rogers et al. 1976). Length was determined from direct measurement of 10 individual of each species.

Biophysical measurements: The effect of ploughing and the addition of legumes and forbs on nitrogen and phosphorus loss in soil water and soil structural properties were measured in Phase 2 on the ‘Typical’ management treatment plots as in Phase 1.

Soil nutrient losses Two teflon sampling tubes were inserted in Phase 1 at random locations within each of the plots under typical management to a depth of 60cm to measure nitrate (NO3) and P concentrations in the drainage water on 3 occasions (November, January and March) during the winter.

Soil chemistry Five soil cores 7.5 cm deep were taken and bulked from each of the plots under typical management in autumn/winter at the start and end of WEB phase 1 and the end of WEB phase 2for soil total carbon, total nitrogen, Olsen extractable phosphorus, total phosphorus and pH analyses using standard approaches (Allen 1974; MAFF 1986).

Soil structure A total of 25 soil penetrometer measurements were made from the surface down to 49cm below the soil surface within the 500 m2 central area of each of the plots under typical management in

2009, 2010 and 2012. This provides a comparative indication of any change in soil surface structure between treatments during the course of the experiment. It should be noted that the degree of resistance shown by soils when tested with penetrometer will be affected by soil moisture. However, calibration of the penetrometer to correct for confounding effects was outside the scope of this study. As penetrometer readings were taken over a short period of time (typically within one to two days within a year) under constant weather conditions differences in soil moisture are not expected to affect between treatment trends. However, between year differences will be confounded to some extent by differences in soil moisture and their interpretation needs to take this into account.

In addition, dry soil bulk density determinations were made. Ten soil cores of 4 cm diameter and 20 cm deep were taken from each typical treatment plot. Each core was sub-divided into a surface 0-10 cm depth section and a 10– 20cm depth section. Each section was weighed fresh and then crumbled and dried to constant weight. Any stones of more than 6mm diameter and any large roots (e.g. tap roots) were removed from a soil section and their total volume measured by volume increase of a known volume of water. The soil bulk density measurements at the two depths provide an indication of whether the establishment of deep rooted species can ameliorate soil structure in the lower soil horizons.

Phospholipid fatty acid analysis (PLFA): Fungal to bacterial ratios Following the methodologies described within De Deyn et al (2011) we determined fungal to bacterial ratios using the Phospholipid fatty acid analysis (PLFA) approach. This approach uses different PLFA markers attributable to either bacterial or fungal origin to assess a ratio of masses of these two groups within the soil. Previous studies suggest that extensification of grassland management is linked to a shift towards the fungal component of the soil, which can be used to infer an increased capacity of the soils to self-regulate their nutrient cycles (Bardgett & McAlister 1999; De Deyn et al 2011).

Statistical analysis Differences in the experimental design undertaken at North Wyke and Jealott’s Hill necessitated different approaches to the analysis presented in subsequent sections.

North Wyke: Repeated measures ANOVA were carried out on data where there were equally spaced time points. The distributions of each variate were checked for normality using probability distribution plots using Genstat version 14 and, where appropriate, normalised using the most appropriate transformations. Residual maximum likelihood (REML) models were used where there were unequally spaced time points. There are some experimental differences between the North Wyke and Jealott’s Hill sites that required modifications. As the North Wyke experiment was not fully factorial when investigating the effects of treatments on response variables several models using various combinations of treatments and sub-sets of treatments were used in the analysis to ensure a fully factorial design. When testing each of the hypothesis shown below an explanation of what model was used to identify significant effects for each hypothesis are specifically highlighted.

Jealott’s Hill: All analyses were undertaken using SAS 9.1 to test the effects of seed mix (Seed), Management (Man.), timing of management (Tim.) cultivation practices (Cult.) and their interaction with the number of years (Year) over which the experiment has been run. The analysis took account of the hierarchical structure of the nesting of treatments within the experimental design (year nested within Cultivation, nested within Time, nested within Man., nested within Seed. Due to this nesting the error term within the ANOVA against which the fixed effects are tested changes according to this hierarchy, whereby Seed is tested against the error term Seed×Block, Management against Seed×Man.×Block, Timing against Seed×Man.×Tim.×Block and Cultivation against

Seed×Man.×Tim.×Cult.×Block. Model simplification is by deletion of least significant factors to form a maximal model containing all interactions of Seed, Man., Tim., Cult. and Year. Deletion continues until the removal of a fixed effect of interaction results in a significant drop in the explained variation of the model. Block is also included as a fixed effect. All counts have been Loge (N+1) transformed to normalise the data. Note that not all analyses use parameters recorded from every experimental plot at Jealott’s hill, for example dry matter yield from cutting by definition did not occur in the grazed plots. In this case this would mean that the fixed effect of management, and all its interaction terms, was not included in the analyses. Where this has occurred this is specifically highlighted.

Part 1. Direct tests of seven key hypotheses.

H1: Ecological generalist legume and non-legume forb species can be readily established in high fertility grassland following conventional or minimal cultivation. Overview: The use of simple seed mixtures sown into existing swards, prepared using either minimum tillage cultivation or deep ploughing seed bed preparation techniques, were successful in introducing the non-legume forb and legume component of the wild flower seed mixtures. Hypothesis 1 was therefore accepted. The success of establishment varied with site, and there was clear evidence that some species were unlikely to be suitable for all soil types. For example, the legumes Melilotus officinalis and Onobrychis viciifolia did poorly in heavy clay soils. Jealott’s Hill, had the highest levels of establishment for both forbs and legumes. Where legumes were sown, Trifolium repens, T. pratense and T. hybridum had summed percentage covers at respectively 20.5 % (± 1.18), 9.54 % (± 0.76) and 6.92 % (± 0.73) of the swards area. With the exception of Cichorium intybus (Asteraceae) (16.6 % ± 0.99), non-legume forbs tended to cover relatively small areas of the sward in the 2009 establishment year of between 0.23 % (± 0.14) for Rumex acetosa (Polygonaceae) and 1.78 % (± 0.23) for Leucanthemum vulgare (Asteraceae). Establishment of legumes and non- legume forbs was also affected by management, with typically cutting being superior to grazing management. H1 was tested in phase 1 and findings were reported in the phase 1 report (BD1466). These results will not be repeated here, but rather we will focus on the interpretation of the results pertaining to hypotheses 2 to 7 in the following sections.

Overview: There was some evidence that the use of minimum tillage or deep ploughing seed bed cultivation practices had an impact on nutrient loss from the soil as part of water leachates. However, this was only the case for Total Phosphorus and not Total Oxidised Nitrogen which showed no response to cultivation method. In contrast to the predictions of hypothesis 2, minimum cultivation methods of seed bed preparation resulted in higher levels of phosphorus loss when compared with deep ploughing. Hypothesis 2 was therefore not only rejected, but the indication is that the predicted impact of cultivation method is reversed.

Jealott’s Hill: Unfortunately the compact clay soils present at Jealott’s Hill proved to be unsuitable for the removal of soil water leachate samples as a result of the extremely low levels of permeability through the soil for water. While the probes were installed in 2010 across all 104 experimental plots the extraction the failure to extract water samples for subsequent analysis meant that this approach had to be abandoned.

Statistical note for North Wyke: To investigate the effects of seed bed preparation on nutrient losses via water leaching, analysis was done on the cultivation method and subsequent management of the sward for the period 2009 – 2012; only one level of seed mix (GLF) was available so this factor was excluded from the analysis. This model therefore identifies the significant effects of all possible interactions between Cultivation,

Management and Year. Mean TON data was log transformed and individual months square root transformed whilst all TP data log transformed to meet the assumptions of the ANOVA.

Soil nutrient loss: Total oxidised nitrogen (TON) North Wyke: No significant effect of seed bed preparation was found on losses of TON in water samples when averaged over the drainage season (November – March) F1,6=0.72, p=0.429 (Fig. 4), with no significant effects of seed bed preparation recorded during the November sampling

(F1,6=3.22, p=0.123), January sampling (F1,6=0.35, p=0.578) or March sampling (F1,6=0.09, p=0.774). Management by cutting was found to have a significant effect on mean TON losses over the whole -1 drainage season compared to grazing (F1,3=24.58, p=0.016; cutting 0.17±0.04 mg l grazing 0.06±0.01 -1 -1 mg l ), though not at the November sampling (F1,3=4.63, p=0.121 cutting 0.20±0.10 mg l grazing -1 -1 -1 0.07 mg l ), January sampling (F1,3=5.26, p=0.106 cutting 0.10±0.02 mg l grazing 0.05±0.01 mg l ) or -1 -1 March sampling (F1,3=3.65, p=0.152 cutting 0.21±0.08 mg l grazing 0.06±0.01 mg l ).

Fig. 4. Effect of cultivation and subsequent management on losses of TON in water samples for 2009-2012 (mean ± SE mean).

Soil nutrient loss: Total phosphorus (TP) North Wyke: A significant effect of seed bed preparation was found on mean TP content in water samples for the whole drainage period (November – March) F1,6=9.70,p=0.021 (minimal cultivation 37.9 ± 11.2 µg l-1 plough 21.5 ± 4.6 µg l-1)(Fig. 5). Cultivation method was significant at the November -1 -1 sampling (F1,6=8.86, p=0.025 minimal cultivation 60.51±21.32 µg l plough 26.23± 6.58 µg l ), and -1 the January sampling (F1,6=6.92, p=0.039 minimal cultivation 21.71±7.47 µg l plough 15.51±3.72 µg -1 -1 l ) though not at the March sampling (F1,6=1.53, p=0.263 minimal cultivation 31.46±10.43 µg l plough 22.72±9.23 µg l-1). No significant effect of management was found on the mean TP losses -1 -1 over the whole drainage period (F1,3=1.79, p=0.273 cutting 30.7 ±11.0 µg l grazing 28.7±5.3 µg l ), however the cutting management produced higher losses of TP compared with the grazing management in the November samples (Man: F1,3=1.45.85, p=0.007). TP content tended to increase throughout the duration of the experiment; TP mean for whole drainage period (Year: F3,36=48.03, p=<0.001), November sampling (Year: F3,36=20.93, p=<0.001), January sampling (Year: F3,36=86.48, p=<0.001) and March sampling (Year: F3,36=65.21, p=<0.001).

Fig. 5. Effect of seed bed preparation and subsequent management of the sward on losses of TP in water samples for 2009- 2012 (mean ± SE mean).

H3: Sown ecological generalist legume and non-legume forb species can regenerate and persist for at least three years under standard/typical grazing or cutting regimes or summer rested regimes.

Overview: As expected, following the autumn sowing in 2008 and subsequent initial successful establishment of legumes and non-legume forbs into the swards, there was a general decline in their percentage covers across all treatments. The only exception to this was in the case of the ‘grass’ only seed mixtures where some natural colonisation of legumes resulted in a small but largely inconsequential increase in their cover. For the non-legume forbs there was a suggestion that maintaining cover after five years was most likely to be achieved under cutting management, particularly if the swards had originally been established using deep ploughing cultivation methods. Independent of management the non-legume forb component was on the whole more persistent within the sward from 2009 to 2012. The loss of legumes was in general far more severe. At one site (North Wyke) the cover of legumes after four full years of monitoring had fallen to levels which rarely exceeded 1-2 %. At the second site (Jealott’s Hill), higher initial establishment seems to have led to better long term persistence of legume cover; particularly where cattle grazing management followed the sowing of the seeds into minimum tillage cultivation was used. It is interesting to note here that grazing was initially the poorer method for establishing legumes, with the highest covers being found where silage cutting management was used. This evidence in part supports hypothesis 3, although best management may be different between legumes and non-legume forbs. Given that legumes are the more sensitive of the two groups, with agricultural cultivars disappearing rapidly, grazing management may be the more effective method overall. As subsequent sections will indicate, management that benefits the long term persistence of legumes does not necessarily translate into the provision of high quality flower forage resources for insect pollinators (where cutting was by far the best approach).

Summed average percentage cover of non-legume forbs

North Wyke: To investigate the effects of management and timing of management in a fully factorial manner on the persistence of non-legume forb species, analysis was done on the management and timing of management treatments and a sub set of the other treatments; only one level of seed mix (GLF) and cultivation (MC) were available, so both of these factors were excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Management, Timing and Year. Non-legume forb data was arcsine transformed to meet the assumptions of the ANOVA. There were no significant management or subsequent timing of management treatment effects on summed non-legume percentage cover. Year did have a significant effect (Year: F3,36=8.14, p=0.003). The cover of non-legume forbs was highest in the establishment year (2009) in the rested treatments but subsequently declined in the following years. In the typical grazed plots, cover of non-legume forbs increased in the second (2010) and third years (2011) and in the typical cut plots cover increased in the second year only. Non-legume forb cover declined in all of the GLF seed mix, minimally cultivated treatments in the final year of the experiment (Fig. 6). 12

6 2009 2010 4

2011

legume forb percentage cover forbpercentage legume - 2 2012

0 MC MC MC MC Summed non Summed GLF GLF GLF GLF Grazed Grazed Cut Cut Typical Rested Typical Rested

Fig. 6. Effect of sward management and the timing of that management on the summed percentage cover of non-legume forbs at North Wyke.

To investigate the effects of management and cultivation in a fully factorial manner on the persistence non-legume forb species, analysis was done on the management and cultivation treatments and a sub set of the other treatments; only one level of seed mix (GLF) and timing of management (T) were available, so both of these were excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Management, Cultivation and Year. Non-legume forb data were arcsine transformed to meet the assumptions of the ANOVA. The percentage cover of non-legume forbs was influenced by the interaction between cultivation and management (Cult. x Man: F1,6=53.05, p=<0.001; Cult: F1,6=240.37, p=<0.001; Man: F1,3=42.63, p=0.007) and the interaction between cultivation and year (Cult. x Year: F3,36=9.93, p=<0.001; Year: F3,36=26.73, p=<0.001). Higher percentage cover of forbs was achieved where ploughing was used to create a seed bed. Cut plots had a significantly higher cover of forbs compared with the grazing management but only in the ploughed treatments. The higher forb cover in the ploughed cut plots can mainly be attributed to chicory (cichorium intybus) which had summed mean cover of 23.5%

(2009), 31.5% (2010), 13.95% (2011) and 6.75% (2012) compared with 4.5% (2009), 10.5% (2010), 3.3% (2011) and 0% (2012) in the ploughed grazed plots. Cover of non-legume forbs tended to be highest in 2010 and then declined in the following two years (Fig. 7). 60

30 2009

legume legume cover forb percentage 2010 - 20 2011

10 2012 Summed nom Summed

0 P MC P MC GLF GLF GLF GLF Grazed Grazed Cut Cut Typical Typical Typical Typical

Fig. 7. Effect of seed bed preparation, sward management and the timing of that management on the summed percentage cover of non-legume forbs at North Wyke.

Jealott’s Hill (summed percentage cover of non-legume forbs): The summed percentage cover of non-legume forbs at Jealott’s Hill responded to an interaction between the establishing seed mixture, sward management and the cultivation method used in initial seed bed preparation in the autumn of 2008 (seed × management × cultivation: F5,42=4.93, p<0.01; Fig. 8). Summed cover of non- legume forbs was perhaps unsurprisingly lowest in the ‘grass’ and ‘grass & legume ‘seed mixes, neither of which had a non-legume forb component in the initial establishing seed mix. The summed percentage cover of forbs in the ‘grass, legume & forb’ seed mix was highest where the plots were cut, with some indication that that deep ploughing in combination with this management promoted the highest covers over the four year period. While grazing management tended to support lower percentage covers of forbs than cutting, the implications of initial seed bed preparation were more significant for grazed than cut plots. Under grazing management the percentage cover of forbs was about c. 45 % lower on average where minimum tillage was used to establish the seed bed compared to conventional deep ploughing.

cover 15

legumeforb percentage Deep - 10 Shallow 5

0 Summednon Cut Graze Cut Graze Cut Graze Grass Grass GL GL GLF GLF Management and seed bed cultivation

Fig. 8. The response of summed non-legume forb percentage cover at Jealott’s Hill to the establishing seed mix, subsequent sward management and the original cultivation method used in seed bed preparation. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

In addition to this, the summed percentage cover of non-legume forbs also had a temporal component, responding to an overall interaction between establishing seed mix, sward management and the timing of this management as well as the years since establishment (seed × management × timing × year: F38,197=10.8, p<0.001; Fig. 9). As seen in the above interaction, the cover of non- legume forbs within the ‘grass’ only and ‘grass & legume’ plots was consistently lower, never exceeding 10 % cover. Focusing on the ‘grass, legume & forb’ swards, the cover of non-legume forbs consistently declined from 2009 to 2012. This decline was most pronounced under cutting management which supported very high cover of forbs in 2009 at c. 50 %, although this fell rapidly in the following year to around c. 20% where it remained. The cover of forbs under grazing management never had this peak in the 2009 establishment year, reaching only 30 % under rested grazing where this management was suspended over the summer. Under rested grazing management the cover of forbs tended to be more persistent over the following years than was seen under cutting, although there was little real difference in cover from 2010 to 2012. The cover of forbs was in general poorest where grazing management had no summer rest period. a) ‘Grass’ seed mix. 60

30 2009

cover 20 2010

legumeforb percentage - 2011 10 2012 0

Summednon Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

b) ‘Grass & Legume’ seed mix

30 2009

cover 20 2010

legumeforb percentage - 2011 10 2012 0

Summednon Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

c) ‘Grass, Legume & Forb’ seed mix. 60

30 2009

cover 20 2010

legumeforb percentage - 2011 10 2012 0

Summednon Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 9. The response of the summed percentage cover of non-legumes forbs at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of its implementation over the four year study period (2009-2012)

Summed average percentage cover of legumes

North Wyke: To investigate the effects of seed mix, management and timing of management in a fully factorial manner on the persistence of legume species, analysis was done on the management and timing of management treatments and a sub set of the other treatments; two levels of seed mix (GL & GLF) but only one level of cultivation (MC) were available, cultivation was therefore excluded from this analysis. This experimental design identifies the significant effects of all possible interactions between Seed Mix, Management, Timing and Year. Legume forb data was arcsine, square root transformed to meet the assumptions of the ANOVA. There were significant interactions between timing of management and year (Timing x Year:

F3,72=4.38, p=0.022; Timing: F1,12=15.06, p=0.022; Year: F3,72=156.22, p=<0.001) on summed percentage cover of legumes at North Wyke. Legume cover was highest in 2009 and then generally declined rapidly over the subsequent years. Legume cover was highest in the typically managed treatments in in both cut and grazed plots compared with the rested after initially being highest in

31 the establishment year (2009) in the rested treatments (Fig. 10). Management and seed mix had no significant effect on legume cover.

40 2009

30 2010 2011 20 2012

10 Summed legume percentage cover percentagelegume Summed

0 MC MC MC MC MC MC MC MC GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Rested Typical Rested Typical Rested Typical Rested

Fig. 10. Effect of seed mix, sward management and the timing of that management on the summed percentage cover of legumes at North Wyke.

To investigate the effects of management and cultivation in a fully factorial manner on the persistence of legume species, analysis was done on the management and cultivation treatments and a sub set of the other treatments; only one level of seed mix (GLF) and timing of management (T) were available, so both of these were excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Management, Cultivation and Year. Legume data were arcsine transformed to meet the assumptions of the ANOVA. There was a significant interaction between the effects of cultivation and year (Cult. x Year:

F3,36=3.64, p=0.049; Cult: F1,6=30.62, p=<0.001; Year: F3,36=87.06, p=<0.001) on summed percentage cover of legumes at North Wyke (Fig. 11). Overall, legume cover was higher were the seed bed had been prepared by ploughing as opposed to minimal cultivation. In 2009 legume establishment was higher in the ploughed cut plots than the minimally cultivated cut plots, however higher cover was found in the grazed minimally cultivated plots compared with the ploughed grazed plots. During the following years ploughing achieved higher legume cover than the minimally cultivated plots under both cut and grazed management. While legume cover declined rapidly from 2009 to 2012 in most of the treatments, in the ploughed grazed plots legume cover showed an increase in 2010 followed by a slight decline in 2011 and then a dramatic decrease in 2012. The percentage cover of legume species was influenced by the interaction between management and year (Man. x Year: F3,36=12.65, p=<0.001; Man: F1,3=0.00, p=0.967) and the interaction between cultivation, management and year (Cult. x Man. x Year: F3,36=4.03, p=0.041). In 2009 under the cutting management legume cover was higher than the grazed management in both the ploughed and minimally cultivated plots. In 2010 and 2011 the grazed ploughed plots generally supported higher legume cover than the cut plots. Legume cover remained higher in the cut minimally cultivated compared with the grazed minimally cultivated plots in the second year of the experiment (2010) however the trend was reversed in 2011 and 2012.

80 70 60 50 40 2009 30 2010 20 2011 10 2012

0 Summed legumepercentage Summedcover P MC P MC GLF GLF GLF GLF Grazed Grazed Cut Cut Typical Typical Typical Typical

Fig. 11. Effect of seed bed preparation, sward management and the timing of that management on the summed percentage cover of legumes at North Wyke.

Jealott’s Hill (Summed percentage cover of legumes): The summed percentage cover of legumes at Jealott’s Hill responded to an interaction between establishing seed mix, sward management, seed bed cultivation method and the number of years since establishment (seed × management × cultivation × year: F33,251=13.4, p<0.001; seed: F2,6=159.5, p<0.001; management: F1,9=2.54, p>0.05; cultivation: F1,66=3.72, p>0.05; year: F3,251=34.8, p<0.001;Fig. 12). Clear differences are apparent between the responses of the three seed mixtures over time. For the ‘grass’ seed mix summed percentage cover of legumes was extremely low in the establishing year of 2009, however the rapid establishment of T. repens meant that by 2012 between 20 -30 % cover was to be found under grazing management. Note, that throughout the course of the experiment legumes became progressively more established into the grass only plots, a factor attributed in part to the dispersal of seeds in cutting machinery as well as seed transfer in the dung of grazing livestock from adjoining plots. To a large part the close proximity of the experimental plots and their relatively small size promoted this process of colonisation into the grass only plots over time. Indeed the establishment of legumes is likely to have occurred at a disproportionately fast rate relative to what would be observed had the grass seed mix been sown into whole fields where the impacts of such seed dispersal mechanisms would be greatly reduced. The establishment of legumes into the grass only plot in the latter years of the study should therefore be interpreted as an artefact of the experimental design which necessitated small plots in close proximity. This temporal response contrasts strongly with that of the ‘grass & legume’ and ‘grass, legume and forb’ seed mixes, both of which show a decline in legume percentage cover over the four year period. For the ‘grass & legume’ seed mix this is really only apparent under cutting management where the percentage cover down from c. 85% in 2009 to between 30-60 % by 2012. Although there is evidence of temporal variation in the percentage cover of legumes under grazing management for the ‘grass & legume’ seed mix, cover tends to also remain at between 30-60 % over the four years. Indeed, for cut plots legume percentage cover by 2012 is the same as that seen in the establishment year 2009 at c. 60 %, although in the intervening years cover did fall below this level. For the ‘grass, legume & forb’ seed mix the trends seen for the ‘grass & legume’ seed mix were repeated to some extent, although under cutting management cover of legumes collapsed over the four years from c. 80 % in 2009 to c. 5 % by 2012. While grazing management initially supported lower legume cover than that of grazed plots (50 – 75% cover), its long term persistence was superior particularly where shallow seed bed cultivation methods where it remained at c. 40 % in 2012.

a) ‘Grass’ seed mix 40 35 30 25 20 2009 15 2010 10 2011 5 2012 Legume sumed percentage cover percentage sumedLegume 0 Deep Shallow Deep Shallow Cut Cut Graze Graze Management and cultivation of seed bed b) ‘Grass & Legume’ seed mix 100 90 80 70 60 50 2009 40 2010 30 2011 20 10 2012 Legume sumed percentage cover percentage sumedLegume 0 Deep Shallow Deep Shallow Cut Cut Graze Graze Management and cultivation of seed bed

c) ‘Grass, Legume & Forb’ seed mix. 90 80 70 60 50 2009 40 2010 30 20 2011 10 2012 Legume sumed percentage cover percentage sumedLegume 0 Deep Shallow Deep Shallow Cut Cut Graze Graze Management and cultivation of seed bed

Fig. 12. The response of the summed percentage cover of legumes at Jealott’s Hill for the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes from 2009 to 2012 in response to sward management and cultivation technique used in seed bed preparation. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth. H4: Establishment of non-legume forbs will incrementally increase the abundance and diversity of pollen and nectar feeding

34 invertebrates compared with grass-dominated or legume rich swards.

Overview: For insect pollinators the inclusion of non-legume forbs within the establishing seed mixtures had an overall positive effect in terms of maintaining both high abundances and species richness within the grassland swards. This effect became more pronounced over time, where the persistence of the non-legume forbs from 2009 to 2012 (see Hypothesis 3 above) helped maintain key foraging resources after the loss of agricultural cultivars of legumes in the initial years. Where rested management was used (either a single early season sward cut or a suspension of grazing over the summer) the increased availability of flowering resources meant that the abundance of foraging pollinators was highest, although this effect was less pronounced that was expected. These general patterns of the need to diversify the floristic composition of the sward were also supported for the beetles, although in terms of both their abundance and species richness the presence of legumes within seed mixes was likely to be as important as the forbs, reflecting the high value of this plant group as food for phytophagous species. It should be noted that for both the pollinators and beetles differences between the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes were in general small in the establishment year (2009), where typically legumes dominated the sward. Overall both hypotheses 4 and 5 were accepted, pointing to the value of a more expensive seed mix containing both legumes and non-legume forbs when establishing the swards if the goals of increased biodiversity and pollination ecosystem service delivery are to be achieved.

Statistical note: The following results focus on the assessment of both hypotheses H4 and H5. Due to structural differences in the experimental design between North Wyke and Jealott’s Hill tests of H4 and H5 were undertaken separately at North Wyke, but in a single combined analysis at Jealott’s Hill. North Wyke H4 and H5 were tested as described below:

H4: To investigate the effects of seed mix in a fully factorial manner on abundance and diversity of pollen and nectar feeding invertebrates at North Wyke, analysis was done on the management treatment and a sub set of the other treatments; only one level of timing of management (typical) and cultivation (MC) were available so both of these were excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed mix, Management and Year.

H5: To investigate the effects of seed mix, management and timing of management in a fully factorial manner on pollen and nectar feeding invertebrates, analysis was done on the seed mix, management and timing of management treatments and a sub set of the cultivation treatment; only one level of cultivation (MC) was available so cultivation was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Timing and Year.

Total pollinator abundance

North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). Total pollinator abundance (honeybees, bumblebees, solitary bees, butterflies and hoverflies) was influenced by an interaction between seed mix and year (Seed x Year: F6,54=5.28, p= 0.001; Seed: F2,6=16.77, p= 0.003; Year: F3,54=7.01, p= 0.002) (Fig. 13). The ‘grass, legume & forb’ seed mix supported the highest numbers of pollinators in the first two years of the experiment compared with the other two seed mixes, however abundance declined over the four years. Total pollinator abundance in the grass only sward was extremely low during the first two years but increased in 2011. During 2012, numbers of pollinators in the grass only sward continued to increase in the cut plots but collapsed completely in the grazed plots. In the ‘grass & legume’ swards the highest pollinator numbers were recorded in the establishment year, abundance declined in 2010 and 2011 followed by an increasing numbers in the final year. Although there was an increase of pollinators in some of the ‘grass only’ and ‘grass & legume’ plots in the final year, abundance was generally low compared with the ‘grass & legume’ and ‘grass, legume & forb’ swards in the cut plots in the establishment year, reflecting an overall temporal trend of declining numbers. Management also had a significant effect on total pollinator abundance (Man. F1,9=13.11, p=0.006) with the cut plots supporting higher numbers than the grazed plots. 25

15 2009 10 2010 2011 5 Total pollinator abundancepollinator Total 2012

0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 13. Effect of seed mix and sward management on total pollinator abundance at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). .

There was a significant interaction between seed mix and year (Seed x Year: F6,108=6.80, p=<0.001; Seed: F2,6=37.87, p=<0.001; Year F3,108=23.95, p=<0.001) on total pollinator abundance at North Wyke (Fig. 14). Peak pollinator numbers were found in the establishment year (2009) in the ‘grass, legume & forb’ sward with slightly lower numbers found in the ‘grass & legume’ seed mix. Pollinator abundance declined in both the ‘grass & legume’ and ‘grass, legume & forb’ swards in 2010 and 2011, however the decline was more rapid in the ‘grass & legume’ sward. In the final year pollinator numbers continued to decline in the ‘grass, legume & forb’ sward however there was a slight increase in numbers from 2011 to 2012 in the ‘grass & legume’ sward. The ‘grass only’ seed mix supported low numbers of pollinators in the establishment year. In later years pollinator abundance did increase in some ‘grass only’ swards, especially in the typically managed plots, however numbers remained relatively low compared with numbers found in the other seed mixtures in the establishment year. The cutting management tended to support more pollinators than the grazing management (Man: F1,9=3.90, p=0.030). Although timing of management did not have a significant

36 effect on pollinator abundance there was an interaction between management and timing of management (Man. x Timing: F1,18=11.95, p=0.003; Timing: F1,18=0.10, p=0.761 ). Cut typically managed plots tended to support higher numbers of pollinators than the cut rested plots whereas the reverse trend was found in the grazed plots with higher pollinator abundance in the rested compared with the typically managed plots. 30

15 2009 2010 10 2011

Total pollinatorabundance Total 2012 5

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Rested Typical Rested Typical Rested Typical Rested Typical Rested Typical Rested

Fig. 14. Effect of seed mix, sward management and timing of management on total pollinator abundance at North Wyke.

Jealott’s Hill (Total pollinator abundance): Combined test of Test of H4 (Establishment of non- legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). Total pollinator abundance (honeybees, bumblebees, solitary bees, butterflies and hoverflies) responded significantly to an interaction between establishing seed mix, management, its timing and the number of years since the implementation of these treatments

(seed × management × timing × year: F33,300=9.38, p<0.001; seed: F2,6=230.8, p<0.001; management: F1,9=33.1, p<0.001; timing: F1,18=28.4, p<0.001; year: F3,300=64.7, p<0.001; Fig. 15). The clearest overall differences in terms of temporal trends are between the three seed mixes, where the ‘grass’ only seed mix tends to have increasing abundance over the four years, while the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes show declining abundance. This trend is perhaps a little deceiving as the abundance of pollinators in the ‘grass’ seed mix are in general low, typically under 7 individuals m-2 in the establishing year compared to 70-90 individual m-2 in the other seed mixes. The underlying cause of this trend reflects the absence of floral resources in the ‘grass’ only seed mix in the establishing year, although as a result of colonisation from surrounding plots by 2012 at least some key floral resource were present and utilised by the pollinators in the ‘grass’ only plots. In contrast, where legumes had been sown into the seed mix there were exceptionally high densities of flowering legumes in the establishment year, particularly under the rested cutting management. These initial high density of legumes followed by the collapse in flower density over the next four years were mirrored by overall pollinator abundance within the experimental plots. Also following

37 the temporal patterns in legume floral resources, plots managed by grazing never supported the same numbers of pollinators as were found under rested cut management (rested cut = c. 70 90 indi. m-2; grazed = c. 30-40 indi. m-2) but at the same time did not show the same level of collapse in pollinator densities. Indeed by 2012 the rested grazed plots supported on average slightly greater densities of pollinators than were found for either rested or typically timings for the cutting management. Where grazing had no rest period over the summer the constant removal of flower heads by feeding cattle meant that floral resources for pollinators were limited. As such pollinator abundance for both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes remained low over the four years under this management, but did not tend to change over this same period. a) ‘Grass’ seed mix 6

3 2009 2010 2

2011 Total pollinator abundance pollinator Total 1 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

b) ‘Grass & Legume’ seed mix 120

100

60 2009 2010 40

2011 Total pollinator abundance pollinator Total 20 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

c) ‘Grass, Legume & Forb’ seed mix

120

100

60 2009 2010 40

2011 Total pollinator abundance pollinator Total 20 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 15. The response of total pollinator abundance at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

Beetle abundance North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). There were significant interaction between management and year (Man. x Year: F3,54=6.86, p=0.004; Man: F1,9=0.19, p=0.0.674; Year: F3,54=50.27, p=<0.001) on beetle abundance at North Wyke (Fig. 16). Beetle numbers increased dramatically in the second year of the experiment and then subsequently declined. Generally more beetles were found in the cut plots than the grazed plots in every year except the establishment year when the reverse trend was found. Significantly more beetles were found in the ‘grass, legume & forb’ sward (Seed: F2,54=17.15, p=0.003) with the ‘grass only’ seed mix supporting the fewest numbers. 100 90 80 70 60 50 2009 40 2010 30 Beetle abumdanceBeetle 20 2011 10 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 16. Effect of seed mix and sward management on beetle abundance at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between timing of management and year (Timing. x

Year: F3,108=12.60, p=<0.001; Timing: F1,18=4.49, p=0.048; Year: F3,108=89.68, p=<0.001) on beetle abundance at North Wyke (Fig. 17). Peak beetle abundance was found in 2010, numbers subsequently declined in the following two years. In the establishment year (2009) beetle numbers

39 were higher in the rested plots than the typically managed plots, however over the next three year period numbers found in the rested and typically managed plots tended to be similar. Overall highest beetle numbers were found in the ‘grass & legume’ sward however numbers found in the

‘grass, legume & forb’ swards were only slightly lower (Seed mix: F2,6=9.98, p=0.012). Although management did not have a significant effect on beetle abundance there was a significant interaction between management and year (Man. x Year: F3,108=3.40, p=0.032; Man: F1,9=0.18, p=0.6.83 ). Grazed plots supported more beetles than the cut plots in the establishment year (2009) however the reverse trend was found for the 2010 to 2012 period especially in 2010. 120

100

60 2009 2010

Beetle abundanceBeetle 2011 40 2012

Fig. 17. Effect of seed mix, sward management and timing of management on beetle abundance at North Wyke

Jealott’s Hill (Beetle abundance): Combined test of Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). The total abundance of sward active beetles collected using suction sampling responded to an interaction between the establishing seed mix, sward management, its timing as well as the number of years since plot establishment (seed × management × timing × year:

F33,252=9.27, p<0.001; seed: F2,6=1053.1, p<0.001; management: F1,9=216.9, p<0.001; timing: F1,18=81.8, p<0.01; year: F3,252=121.9, p<0.001; Fig. 18). In parallel to the responses seen for the pollinators, the abundance of beetles within the sward was dramatically higher where legumes were part of the establishing seed mix (‘grass & legume’ and ‘grass, legume & forb’ seed mixes). This initial high abundance was most pronounced in 2009, as large numbers of Protapion spp. seed weevils (Coleoptera: Apionidae) demonstrated a strong numeric response to the cover of clovers. For this reason c. 13 – 18 times more beetles (principally made up of these weevils) were found in

2009 for the rested cutting management of the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes that for comparable treatments of the ‘grass’ only seed mix. The subsequent collapse in legume flower densities across all treatments was mirrored by the collapse in beetle abundance, particularly between 2009 and 2010. a) ‘Grass’ seed mix 30

15 2009 Beetleabundance 2010 10 2011 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing b) ‘Grass & Legume’ seed mix 450 400 350 300 250 2009

Beetleabundance 200 2010 150 100 2011 50 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

c) ‘Grass, Legume & Forb’ seed mix

600

500

400

300 2009 Beetleabundance 2010 200 2011 100 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 18. The response of beetle abundance at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

In addition to the above higher order interaction, beetle abundance was significantly affected by an interaction between the establishing seed mix and the cultivation practice used in initial seed bed preparation (seed × cultivation: F2,45=4.35, p=0.03; cultivation: F1,45=7.51, p<0.01; Fig. 19). Although the general trend of peak abundance being found in the establishment year is maintained, the use of shallow cultivation to establish the seed bed was seen to benefit beetle abundance in this year. It is at least potentially possible that shallow cultivation may have caused less damage to beetles overwintering within the sward that deep ploughing, and that the benefits of this effect became relatively unimportant in subsequent years where such seed bed preparation was not applied. Perhaps more likely is that this response reflects impacts on legume cover in the establishment year. However, while the summed percentage cover of legume was highest under shallow cultivation where grazing management was applied, this trend was reversed (albeit only slightly) for cutting management. It is likely that the establishment of some legume species were of greatest benefit in terms of host plants for seed weevils did best under shallow cultivation, for example red clover.

250

200

150 2009 2010 100

2011 Beetleabundance 50 2012

0 Deep Shallow Cultivation

Fig. 19. The response of beetle abundance at Jealott’s Hill in each of the three establishing seed mixtures to the cultivation methods used in seed bed preparation in 2008. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Bumblebee abundance North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). There was a significant interaction between seed mix and year (Seed x Year: F6,54=12.04, p=<0.001; Seed: F2,6=38.82, p=<0.001; Year F3,54=34.74, p=<0.001) on bumblebee abundance at North Wyke (Fig. 20). Generally bumblebee abundance declined throughout the experiment. In the establishment year the ‘grass, legume & forb’ seed mix supported the highest numbers of bumblebees with the grass only sward supporting the fewest numbers. The ‘grass & legume’ and ‘grass, legume & forb’ swards continued to support more bumblebees than the ‘grass only’ sward in 2010, however the ‘grass only’ sward supported the most bumblebees in 2011 although numbers were low. There was also a significant interaction between management and year (Man. x Year: F3,54=4.29, p=<0.019; Man: F1,9=23.11, p=<0.001) with higher bumblebee abundance in the cut plots compared with the grazed plots in all years. No bumblebees were recorded at all in the final year. 20 18 16 14 12 10 2009 8 2010 6

4 2011 Bumblebeeabundance 2 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 20. Effect of seed mix and sward management on bumblebee abundance at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between seed mix and year (Seed x Year: F6,108=16.37, p=<0.001; Seed: F2,6=37.04, p=<0.001; Year F3,108=78.14, p=<0.001) on bumblebee abundance at North Wyke (Fig. 21). Peak bumblebee abundance was found in the establishment year and numbers subsequently declined throughout the experiment with no bumble bees recorded in the final year. The ‘grass, legume & forb’ sward supported the highest number of bumblebees and the ‘grass only’ sward supported the least. Cut plots supported higher bumblebee abundance than grazed plots (Man: F1,9=12.45, p=0.005). Although the timing of management was not significant there was an interaction between timing and year (Timing. x Year: F3,108=4.34, p=0.014; Timing: F1,18=0.07, p=0.793) and timing and management (Timing. x Man: F1,18=9.05, p=0.008). Overall, in 2009 the rested plots supported higher bumblebee densities than the typically managed plots with the reverse trend in 2010 and 2011 however there was a difference between the two sward managements. The cut typically managed plots tended to support more bumblebees than the cut rested plots whereas the reverse trend was found under the grazing management.

10 2009

8 2010 2011

Bumblebeeabumdance 6 2012 4

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut TypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRested

Fig. 21. Effect of seed mix, sward management and timing of management on bumblebee abundance at North Wyke

Jealott’s Hill (Bumblebee abundance): Combined test of Test of H4 (Establishment of non- legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between establishing seed mix, sward management, the timing of management and years since plot establishment on the abundance of bumblebees at Jealott’s Hill (F40, 190=8.06, p<0.001). The clearest overall trend was the effect of seed mix, with the ‘grass’ only seed mix supporting almost no bumblebees relative to the two seed mixtures containing wild flowers (Fig. 22). In the case of the ‘grass & legume’ and ‘grass, legume & forb seed mixes’ there was a consistent trend of decreasing bumblebees abundance over time that was associated with all combinations of sward management and it timing. The only exception to this was a slight overall depression in bumblebee abundance in 2010, although this was linked with effects of a drought year. For both of the floristically diverse seed mixtures higher abundances of bumblebees were supported under cutting management. Independent of management type, a cessation of either cutting or grazing over the summer months under rested management supported higher abundance of bumblebees. This is linked with the phenological development of flowers heads within the ‘grass & legume’ and ‘grass, legume & forb’ seed mixtures where management was suspended. Overall, it seems that slightly higher abundances of bumblebees were linked with the diverse ‘grass, legume & forb’ seed mix. There was no significant effect of seed bed cultivation, either on its own or as part of an interaction (p>0.05).

44 a) ‘Grass’ seed mix 80 2009 60 2010 2011 40 2012

20 Bumblebee Bumblebee abundance 0 Rest Typ. Rest. Typ. Cut Cut Graz. Graz.

Sward management and its timing b) ‘Grass & Legume’ seed mix 80 2009 60 2010 2011 40 2012

20 Bumblebee Bumblebee abundance 0 Rest Typ. Rest. Typ. Cut Cut Graz. Graz.

Sward management and its timing c) ‘Grass, Legume & non-legume forb’ seed mix 80 2009 60 2010 2011 40 2012

20 Bumblebee Bumblebee abundance 0 Rest Typ. Rest. Typ. Cut Cut Graz. Graz.

Sward management and its timing

Fig. 22. The response of bumblebee abundance at Jealott’s Hill for the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

Butterfly abundance

North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). Butterfly abundance responded significantly to management (Man: F1,9=8.19, p=0.019) with the cutting regime supporting more butterflies than the grazing regime (Fig. 23). Butterfly numbers were low in all plots and there were no significant seed mix or year effects. 8 7 6 5 4 2009 3 2010 2

Butterfly abundanceButterfly 2011 1 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 23. Effect of seed mix and sward management on butterfly abundance at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). The ‘grass, legume & forb’ swards supported the highest butterfly numbers and the ‘grass only’ sward supported the least (Seed: (F2,6=19.88, p=0.002). Butterfly abundance was influenced by management (F1,9=5.53, p=0.043) and year (F3,108=4.58, p=0.014) (Fig. 24). Butterfly numbers tended to be higher in the cut plots compared with the grazed plots. There were no clear temporal patterns with butterfly numbers increasing over time in some treatments and decreasing in others, however overall densities were highest in the establishment year (2009) and 2011 with very low numbers recorded in the final year. No significant timing of management effects were found.

4 2009 2010 3

Butterfly abundance Butterfly 2011

2 2012

Fig. 24. Effect of seed mix, sward management and timing of management on butterfly abundance at North Wyke

Jealott’s Hill (Butterfly abundance): Combined test of Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between establishing seed mix, sward management, the timing of management and the number of years since plot establishment on the abundance of butterflies found at Jealott’s Hill (F40, 190=2.78, p<0.001; Fig. 25). In contrast to most of the other pollinator groups, seed mix was of limited importance in dictating the number of butterflies encountered within the plots. Although the abundance of butterflies was slightly higher in the ‘grass & legume’ and ‘grass, legume & forb’ plots, it was typically low across all seed mixes rarely exceeding an average observation of 4 individuals per transect in the three year period from 2010 to 2012. Typically the majority of this abundance was made up of observations of the Meadow Brown (Maniola jurtina), which with grass feeding larvae may have been utilising any of the plots for sites to lay eggs in rather that to forage as adults for nectar on flowers. The clearest exception to this was in the establishment year of 2009, where very high abundances were encountered in the rested cut plots. This was linked with a particularly large migration of the Painted Lady butterfly (Vanessa cardui) in that year (http://www.ukbutterflies.co.uk/species.php?species=cardui). There was a clear preference by this species for the sward that established following the first cut for the ‘grass & legume’ and ‘grass, legume & forb’ plots. It is thought that the rapid recovery of the flowering component of the sward in 2009 following the first sward cut coincided with the normal peak abundance of this species from July to September. Under grazing management the more continuous removal of vegetation reduced the utility of these plots as a foraging habitat for the Painted Ladies. Outside of what is arguably an anomalous year, the abundance of butterflies showed only minimal differences between the different seed mixes, management types and management intensities. In addition, there was no significant effect of seed bed cultivation practice

47 as implemented in 2008 to establish a seed bed, either on its own or as part of a significant interaction (p>0.05). a) ‘Grass’ seed mix 12

10 2009 2010 8 2011 6 2012

Butterfly abundance Butterfly 2

0 Rest Typ. Rest. Typ. Cut Cut Graz. Graz.

Sward management and its timing b) ‘Grass & Legume’ seed mix 12

10 2009 2010 8 2011 6 2012

Butterfly abundance Butterfly 2

0 Rest Typ. Rest. Typ. Cut Cut Graz. Graz.

Sward management and its timing c) ‘Grass, Legume & non-legume forb’ seed mix 12

10 2009 2010 8 2011 6 2012

Butterfly abundance Butterfly 2

0 Rest Typ. Rest. Typ. Cut Cut Graz. Graz.

Sward management and its timing

Fig. 25. The response of butterfly abundance at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

Honeybee abundance

2 2011 Honeybee abundanceHoneybee 1 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 26. Effect of seed mix and sward management on honeybee abundance at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between seed mix and year (Seed mix x Year:

F6,108=4.79, p=<0.001; Seed mix: F2,6=18.94, p=0.003; Year F3,108=7.96, p=<0.001) on honeybee abundance at North Wyke (Fig. 27). Highest honeybee densities were found in the establishment year (2009) followed by the final year (2012) with the ‘grass & legume’ sward supporting the highest numbers and the ‘grass only’ sward the least (0 numbers) in both of these years. In 2010 the ‘grass, legume & forb’ seed mix supported the highest honeybee densities and then subsequently declined in the final two years. No significant management or timing of management effects were found on honeybee abundance but there was a significant interaction between these two treatments (Man. x

Timing: F1,18=6.68, p=0.019; Man: F1,9=0.09, p=0.771; Timing F1,18=0.33, p=0.574). The grazed rested plots tended to have higher honeybee numbers compared with the grazed typically managed plots with the reverse trend found under the cutting management.

4 2009 2010 3

2011 Honeybee abundanceHoneybee 2 2012

Fig. 27. Effect of seed mix, sward management and timing of management on honeybee abundance at North Wyke

Jealott’s Hill (Honeybee abundance): Combined test of Test of H4 (Establishment of non- legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). Honeybee abundance responded significantly to a higher order interaction between establishing seed mix, management and its timing and the number of years since the implementation of these treatments (seed × management × timing × year: F33,300=8.85, p<0.001; seed: F2,6=213.1, p<0.001; management: F1,9=56.6, p<0.001; timing: F1,18=20.5, p<0.001; year: F3,300=48.4, p<0.001; Fig. 28). As for total pollinator abundance, the ‘grass’ only seed mix provided few floral resources and as a result typically supported extremely low abundances of honeybees, equivalent to fewer than 1 individual per transect walk. For both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes honeybee abundances were considerably higher in the establishment year (equivalent to 20-30 per transect walk). Again honeybee abundances followed the general pattern described for the overall pollinators, in that it tracked the initially high availability of legume flowers under the rested cutting management and followed the collapse of this resource over the four year period to 2012. The unexpected peak in honeybee abundance in the ‘grass & legume’ seed mix in 2011 corresponds to a similar, although unexplained spike in non- legume forb flower density in the same year. In general cutting management, in particular rested management (single yearly cut), supported the highest abundances of bees during the establishment years. However, after four years the initially lower abundances of honeybees found under the rested grazing management were maintained to a greater extent and so were higher than that found for the rested cut plots at least for the ‘grass, legume & forb’ seed mix. Indeed the better overall performance for the four year life of the experimental plots in the ‘grass, legume & forb’ seed mix is attributed to the rise in availability of non-legume forb flowers which counteracted the collapse in density of legume flowers over the same period.

50 a) ‘Grass’ seed mix 1.4

1.2

0.8 2009 0.6

2010 Honeybeeabundance 0.4 2011 0.2 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing b) ‘Grass & Legume’ seed mix 30

15 2009 2010

Honeybeeabundance 10 2011 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing c) ‘Grass, Legume & Forb’ seed mix 35

20 2009 15

2010 Honeybeeabundance 10 2011 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 28. The response of honeybee abundance at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

Hoverfly abundance

North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). Hoverfly abundance was influenced by management (F1,9=5.34, p=0.046) with higher numbers generally found in the cut plots than the grazed plots (Fig. 29). No significant seed mix or year effects were found. 12

6 2009

4 2010

Hoverfly abundance Hoverfly 2011 2 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 29. Effect of seed mix and sward management on hoverfly abundance at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). No significant temporal effect on hoverfly abundance was found but there was a significant interaction between seed mix and year (Seed x Year: F6,108=2.63, p=0.025; Seed: F2,6=3.56, p=0.040; Year F3,108=2.07, p=0.115) (Fig. 30). Lowest hoverfly numbers were recorded in the establishment year (2009) in all sward mixtures. In 2009 and 2010 the highest hoverfly densities were found in the ‘grass, legume & forb’ sward, numbers subsequently declined in this sward type in the following years. Hoverfly abundance remained low in the ‘grass & legume’ sward and in the ‘grass only’ swards numbers tended to increase in 2011 and 2012 however densities remained low. No significant management or timing of management effects were found on hoverfly abundance but there was a significant interaction between these two treatments (Man. x Timing: F1,18=15.65, p=<0.001; Man: F1,9=0.02, p=0.904; Timing: F1,18=1.59, p=0.223). The grazed rested plots tended to have higher hoverfly numbers compared with the grazed typically managed plots with the reverse trend found under the cutting management.

6 2009 2010

Hoverfly abundance Hoverfly 4 2011 2012

Fig. 30. Effect of seed mix, sward management and timing of management on hoverfly abundance at North Wyke

Jealott’s Hill (Hoverfly abundance) : Combined test of Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). The abundance of hoverflies (Syrphidae) responded to a higher order interaction between establishing seed mix, management and its timing as well as the number of years since the implementation of these treatments (seed × management × timing × year: F33,252=8.84, p<0.001; seed: F2,6=158.2, p<0.001; management: F1,9=34.8, p<0.001; timing: F1,18=41.2, p<0.001; year: F3,252=56.3, p<0.001; Fig. 31). As for previously described responses for pollinator abundance, the absence of a sown component of floral resources (legumes of non-legume forbs) in the ‘grass’ only seed mix meant that the value of these plots was almost insignificant for foraging hoverflies (c. 1 individual per transect walk). For hoverflies, which in contrast to bumblebees forage to a larger extent on non-legume forbs, the ‘grass, legume & forb’ seed mix clearly supported the highest densities over the four years. This is directly linked to the increasing densities of the non-legume forbs sown into this seed mix the flowering resources of which increased from 2009-2012. This increase in floral resources was mirrored by the hoverflies, so that by 2012 cut or rested grazed plots supported higher densities that in 2009. For the ‘grass & legume’ seed mix, the absence of this non- legume forb resource meant that while the high legume densities in the establishment year did benefit hoverflies (particularly in the rested cut treatment), the subsequent collapse in this resource in following years meant that this seed mix was of limited long-term value for hoverflies.

53 a) ‘Grass’ seed mix 0.8 0.7 0.6 0.5 0.4 2009

Hoverfly abundance Hoverfly 0.3 2010 0.2 2011 0.1 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing b) ‘Grass & Legume’ seed mix 16 14 12 10 8 2009

Hoverfly abundance Hoverfly 6 2010 4 2011 2 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing c) ‘Grass, Legume & Forb’ seed mix 30

15 2009

Hoverfly abundance Hoverfly 2010 10 2011 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 31. The response of hoverfly abundance at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

There was also a significant interaction between establishing seed mix and the cultivation practice used in the initial preparation of the seed bed (seed × cultivation: F2,45=3.18, p=0.05; cultivation: F1,45=9.82, p<0.01; Fig. 32). This significant effect is likely to be underpinned by flower density for

54 the non-legume forbs (see above) which was highest where deep cultivation had been used to create a seed bed for the ‘grass, legume & forb’ seed mix. Again this clearly points to the importance of non-legume forbs for at least a proportion of the pollinator community as a whole. 14

6 Deep Shallow

4 Hoverfly abundance Hoverfly

0 Grass GL GLF Seed mix

Fig. 32. The response of hoverfly abundance at Hill to the establishing seed mixture and the cultivation practice used to establish its seed bed in 2008.

Beetle species richness

North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). There was a significant interaction between management and year (Man. x Year: F3,54=7.74, p=<0.001; Man: F1,9=0.39, p=0.545; Year F3,54=31.83, p=<0.001) on beetle species richness (Fig. 33). In the establishment year beetle diversity was higher in the grazed plots than the cut plots, however the reverse trend was found in all the other years. Peak beetle abundance was recorded in the second year of the experiment (2010) with the second highest diversity found in 2011 in most instances. Beetle abundance was influenced by seed mix (F2,6=10.60, p=0.011) with higher diversity found in the ‘grass & legume’ seed mix and lowest in the ‘grass only’ seed mix. 14 12 10 8 2009 6 2010 4 2011 Beetle speciesBeetlerichness 2 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 33. Effect of seed mix and sward management on beetle species richness at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between timing of management and year (Timing x

Year: F3,108=9.18, p=<0.001; Timing: F1,18=18.52, p=<0.001; Year F3,108=50.56, p=<0.001) on beetle species richness at North Wyke (Fig. 34). Beetle diversity increased in the second year of the experiment and then subsequently declined. Rested plots supported higher beetle diversity than the typically managed plots although the differences were not as marked in the final two years.

Seed mix also influenced beetle species richness (Seed: F2,6=21.95, p=0.002) with higher diversity found in the ‘grass & legume’ seed mix and lowest in the ‘grass only’ seed mix. Management had no significant effect however there was a significant interaction between management, timing of management and year (Man. x Timing x Year: F3,108=4.75 p=0.009; Man: F1,9=0.03 p=0.859). The rested grazed plots supported higher beetle diversity in all years than the typically grazed plots however this was only the case in the cut rested plots in years 2009 and 2012.

10 2009 8 2010

6 2011 Beetle species richnessspecies Beetle 2012 4

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut TypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRested

Fig. 34. Effect of seed mix, sward management and timing of management on beetle species richness at North Wyke

Jealott’s Hill (Beetle species richness): Combined test of Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). Total beetle species richness responded significantly to establishing seed mix, sward management, its timing as well as the number of years since plot establishment (seed × management × timing × year: F33,300=4.14, p<0.001; seed: F2,6=496.8, p<0.001; management: F1,9=20.4, p<0.001; timing: F1,18=71.5, p<0.01; year: F3,300=50.5, p<0.001; Fig. 35). The most significant overall response was to establishing seed mix, with the inclusion of legumes being crucial if higher diversities of beetles are to be supported. In general the grass only seed mix typically

56 supported between 3 – 8 species of beetle, although there was a higher peak (c. 10-12 species) under rested management during the establishment year. In general, this diversity was made up of generalist predatory / polyphagous species, in particular rove and ground beetles. The significantly higher species richness of beetles found in association with the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes is linked with the colonisation by phytophagous beetles utilising legumes. For both ‘grass & legume’ and ‘grass, legume & forb’ seed mixes, rested management (single sward cut or suspended summer grazing) supported the highest diversities. This is likely a product of reduced management pressure which allowed the phenological development of plant structures fed upon by beetle larvae, e.g. seed heads, flower heads and to some extent stems (Morris, 2000; Woodcock et al., 2009; Woodcock et al., 2007). Consistent with the collapse in both cover and species richness of legumes throughout the four year succession, beetle species richness also declined consistently across all plots from 2009 to 2012. For both ‘grass & legume’ and ‘grass, legume & forb’ seed mixes this was equivalent to a loss of c. 10 species from plots receiving rested cutting or grazing management. This loss of species richness is linked not only to an overall decline in legume cover and species richness, but also to the loss of specific plant species that support a disproportionately high diversity of phytophagous beetles, e.g. red clover is fed on by many seed feeding weevils (Bullock, 1992). Fig. 36 shows the disproportionate importance of legumes for plant feeding beetles within the Jealott’s hill site (Woodcock et al 2012). In particular, the clovers are important for phytophagous beetles with red clover supporting the greatest number of feeding species, although white clover is an important host plant. This importance of legumes is most apparent for the ‘rested grazed ‘grass & legume’ seed mix which supported beetle species richness at high levels (c. 20 species) from 2009-2011, but showed a collapse in 2012 as red clover and several other legume species were almost entirely lost. In the context of these floristically diversified ex- improved grasslands it appears that the non-legume forb component of the ‘grass, legume & forb’ seed mix is of limited importance in supporting high diversities of beetles. This is in contrast to the pattern seen for some pollinators. For examples, in the case of the hoverflies the prevalence of non-legume forbs in the latter years of the succession for the ‘grass, legume & forb’ plots were crucial in maintaining high numbers by 2012. It appears that non-legume forbs are of comparatively little importance for many phytophagous beetles, either in terms of abundance or species richness. This is not to say that there were not specific beetle associations with non-legume forbs, only that their inclusion in the seed mix failed to result in large scale increases in species richness. a) ‘Grass’ seed mix 16 14 12 10 8 2009 6 2010 Beetle species richness Beetle species 4 2011 2 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

b) ‘Grass & Legume’ seed mix 25

15 2009 10 2010 Beetle species richness Beetle species 2011 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing c) ‘Grass, Legume & Forb’ seed mix 30

15 2009 2010 10 Beetle species richness Beetle species 2011 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 35. The response of beetle species richness at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

a) ‘Grass’ seed mix

Is.vi Pr.di Pr.tr Si.hi Si.le Si.li Ty.pi Ag.li 6 7 8 3

Tr.re Tr.pr Gr. Lo.co Pl.ma

b) ‘Grass & legume’ seed mix Ty.pi Hy.pu Pr.as 8 Si.li Hy.ni Si.le Pr.di Si.hi Is.vi Pr.tr 9,10, 2,11,12,13

Lo.co Tr. hy Tr.re Tr.pr Ci.ar Ru.ac

c ) ‘Grass, legume & non-legume forb’ seed mix Pr.as Hy.ni Hy.pu 8, 14, 9 Si.li Si.hi Ty.pi Si.le Pr.di Is.vi Pr.tr 10, 2, 3, 4, 6, 7, 13, 5

Lo.co Me.lu. Tr.du Tr.hy Tr.re Tr.pr Ci.ar Gr. Pl.ma Ru.ac BEETLES: Si.li.=Sitona lineatus; Si.hi.=S. hispidulus; Si.le.=Sitona lepidius; Ty.pi.=Tychius picirostirs; Is.vi=Ischnopterapion virens; Pr.di= Protapion dichroum; Pr.as=P. assimilie; Pr.tr.=P. trifolii; Hy.ve.=Hypera venusta; Hy.ni.=H. nigrirostris; Hy.pu.=H. punctata; Ag.li.=Agriotes lineatus; 1=Tychius picirostris; 2=Asiorestia ferruginea; 3=Longitarsus luridus; 4=Longitarsus pratensis; 5=Perapion marchicum; 6=Chaetocnema hortensis; 7=Oulema melanopa; 8=Ischnopterapion loti; 9=Sitona sulcifrons; 10=Protapion apricans; 11=Ceratapion cardorum; 12=Sphaeroderma rubidum; 13=Chaetocnema concina; 14=Tychius stephensi. PLANTS: Lo.co.=Lotus corniculatus; Tr.=Trifolium (du.= dubium, hy = hybridum; re=repens, pr=pratense); Me.lu = Medicago lupulina; Ci.ar=Cirsium arvense; Pl.ma=Plantago major; Ru.ac=Rumex acetosa.; Gr.=all grass species.

Fig. 36. This figure shows the increased complexity of trophic interactions (summed from 2009 – 2011) between plant feeding beetles within the three seed mixtures and their host plants at Jealott’s Hill (reproduced from Woodcock et al (2012)). Further detail of the methodology is given in this paper, but in summary individual beetle species are represented by bars on the upper tier, the length of which is proportional to the summed biomass of that beetle species feeding on that plant. Individual plants are represented by bars on the lower tier, the length of these bars is proportional

59 to the total biomass of beetles with feeding associations with those plants. Species abbreviations refer to the first and second letters of the generic and specific names, or numbers where space is limited (see caption below figure). Only food webs under intensive grazing management are shown for the i) grass only seed mix, iii) grass & legume seed mix, and iv) grass, legume and forb seed mix.

Bumblebee species richness

North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). There was a significant interaction between seed mix and year (Seed mix. x Year: F6,54=11.68, p=<0.001; Seed: F2,6=16.17, p=0.004; Year F3,54=30.08, p=<0.001) on bumblebee species richness at North Wyke (Fig. 37). Overall the highest bumblebee diversity was found in the ‘grass, legume & forb’ sward. The ‘grass & legume’ and ‘grass, legume & forb’ swards supported comparable bumblebee species richness in the establishment year (2009) whereas unsurprisingly in the ‘grass only’ swards diversity was low. In 2010 bumblebee diversity completely collapsed the ‘grass & legume’ seed mix whereas diversity was more stable and increased in the ‘grass, legume & forb’ sward. Bumblebee species richness was low in all of the seed mixtures in 2011 and during the final year no bumblebees were recorded in any of the plots. Bumblebee diversity was greater in the cut plots than the grazed plots (Man: F1,9=11.20, p=0.009). 4 3.5 3 2.5 2 2009 1.5 2010 1 2011 0.5 Bumblebeespeciesrichness 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 37. Effect of seed mix and sward management on bumblebee species richness at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). There was a significant interaction between seed mix, timing of management and year

(Seed x Timing x Year: F6,108=3.66, p=0.013; Seed: F2,6=71.15, p=<0.001; Timing: F1,108=8.13, p=0.011; Year F3,108=64.32, p=<0.001) on bumblebee species richness at North Wyke. Bumblebee diversity was overall highest in the establishment year (2009) with a few treatments supporting the highest densities in 2010 (Fig. 38), diversity then declined in the following two years. The ‘grass, legume & forb’ seed mix supported the highest number of bumblebee species over the four year period (Seed mix x year: F6,108=9.84, p=<0.001) and the ‘grass only’ seed mix supported the least. The rested plots tended to support more bumblebee species than the typically managed plots in all years (Timing x year: F3,108=3.97, p=0.028) except the ‘grass, legume & forb’ sward in 2010 and all sward types in 2011 however differences were negligible as numbers of species were extremely low. Management also influenced bumblebee species richness (Man: F1,9=25.22, p=<0.001) with more bumblebee species found in the cut plots than the grazed plots.

4.5

3.5

2.5 2009 2010 2 2011

1.5 2012 Bumblebee speciesrichness Bumblebee

0.5

0 MC MC MC MC G G G MC G MC Cut MC MC Cut GL GL MC GL GL MC MC GLF Cut GLF GLF GLF Cut Grazed Grazed Typical Typical GrazedRestedRested Grazed Cut Typical Grazed Rested Rested Typical Grazed Cut Typical Rested Rested Typical

Fig. 38. Effect of seed mix, sward management and timing of management on bumblebee species richness at North Wyke

Jealott’s Hill (Bumblebee species richness): Combined test of Test of H4 (Establishment of non- legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). Bumblebee species richness differed significantly between seed mixtures in response to an interaction with the number of years since establishment (seed × year:

F2,383=17.5, p<0.001; seed: F2,6=144.3, p<0.001; year: F3,383=21.3, p<0.001; Fig. 39). Many bumblebee species actively utilise legumes as a principal floral foraging resource, and so the general pattern in species richness seems to reflect trends observed in the density of this flowering resource as already described above. Highest legume flower density was found in the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes in the establishment year (2009), with this collapsing thereafter. This general pattern was also seen for the bumblebees with peak species richness linked with seed mixes containing legumes in the establishing year. While the species richness of bumblebees was comparable between the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes in 2009, it collapsed at a much greater rate in the former of these (2009 = c. 5 species; 2012 = c. 2 species). For the ‘grass, legume & forb’ seed mix the species richness of bumblebees supported was far more robust, losing on average only one species per plot over this period (2009 = c. 5 species; 2012 = c. 4 species). It is likely this reflects the maintenance of floral resources in the ‘grass, legume & forb’ seed mix, which while progressively losing legume flowers over the four year period was supplemented by a rapid increase in non-legume forb flowers. Independent of these overall differences resulting from the addition of non-legume forbs to legumes within the seed mix, the absence of both of these in the ‘grass’ only seed mix meant that species richness of bumblebees was consistently low (c. 1 species).

4 2009 3 2010 2 2011 2012

1 Bumblebee species richness Bumblebee species

0 Grass GL GLF Seed mix

Fig. 39. The response of bumblebee species richness to the establishing seed mixture from 2009 to 2012 at Jealott’s Hill.

There was also a significant higher order interaction between establishing seed mix, management and the timing with which that management was applied (seed × management × timing: F5,18=3.80, p=0.02; management: F1,9=6.84, p=0.03; timing: F1,18=5.60, p=0.03; Fig. 40). Following the trends described above bumblebee species richness was highest in seed mixes containing either legumes or non-legume forbs. In addition there was some evidence that species richness was on average slightly higher (c. 0.25 – 0.5 species) where rested management (one sward cut or no summer grazing) was used.

5 4.5 4 3.5 3 2.5 2 Rested 1.5 Typical 1

Bumblebee species richness Bumblebee species 0.5 0 Cut Graze Cut Graze Cut Graze Grass Grass GL GL GLF GLF Management and its timing

Fig. 40. The response of bumblebee species richness at Jealott’s Hill to the establishing seed mixture and the timing and type of management subsequently applied to its sward.

Finally, the cultivation practice used to establish the seed bed in 2008 had a significant effect on bumblebee species richness (F1,47=5.54, p=0.02; Fig. 41). Overall deep cultivated plots tended to support fewer species of bumblebee that occurred where shallow cultivation had been used to establish the seed bed. We have no direct mechanistic explanation for this, as legume flower

62 density was not influenced by seed bed cultivation practice, while deep cultivation promoted the density of non-legume forb flower cover.

2.5

1.5

0.5 Bumblebee species richness Bumblebee species

0 Deep Shallow Cultivation

Fig. 41. The response of bumblebee species richness at Jealott’s Hill to the cultivation practice used during initial seed bed preparation in 2008. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Butterfly species richness

North Wyke: Test of H4 (Establishment of non-legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards). Management significantly affected butterfly diversity (Man: F1,9=9.98, p=0.012) with more species found in the cut plots than the grazed especially in the ‘grass & legume’ and ‘grass, legume & forb’ swards (Fig. 42). Year and seed mix had no significant effect on butterfly species richness. 2 1.8 1.6 1.4 1.2 1 2009 0.8 2010 0.6 0.4 2011 Butterfly speciesButterflyrichness 0.2 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF Grazed Cut Grazed Cut Grazed Cut Typical Typical Typical Typical Typical Typical

Fig. 42. Effect of seed mix and sward management on butterfly species richness at North Wyke.

Test of H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). Butterfly abundance was influenced by management (F1,9=4.96, p=0.049) and year (F3,108=6.58, p=0.003)(Fig. 43). Butterfly numbers tended to be higher in the cut plots compared with

63 the grazed plots. There were no clear temporal patterns however overall species richness was highest in the establishment year with very low numbers of butterfly species recorded in the final year. The more diverse swards, ‘grass, legume & forb’, followed by the ‘grass & legume’ seed mixes supported the most butterfly species (Seed: F2,6=6.26, p=0.034). No significant timing of management effects were found. 2.5

1.5 2009 2010 1 2011

2012 Butterfly speciesButterflyrichness

0.5

Fig. 43. Effect of seed mix, sward management and timing of management on butterfly species richness at North Wyke

Jealott’s Hill (Butterfly species richness): Combined test of Test of H4 (Establishment of non- legume forbs will be of greater benefit to invertebrates compared with grass-dominated or legume rich swards) and H5 (Rested management will benefit invertebrates of legume rich or legume and forb rich swards). Butterfly species richness responded to a significant higher order interaction between establishing seed mix, sward management, the timing of its application and overall sampling year (seed × management × timing × year: F33,300=1.73, p<0.01; seed: F2,6=10.3, p<0.05; management: F1,9=0.71, p>0.05; timing: F1,18=8.43, p<0.01; year: F3,300=15.1, p<0.001; Fig. 44). However, butterfly species richness was on average extremely low in all plots, and rarely exceeded an average of 2.5 species, one of which was typically the ubiquitous meadow brown (Maniola jurtina L.). For this reason there was little evidence of the experimental plots representing a key resource for butterflies. The ‘grass’ only seed mix typically supported the lowest species richness of butterflies. In both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes higher butterfly species richness was found in the rested cut plots, although this occurred only in the establishing year (2009). These was no significant effect of initial seed bed cultivation, either as an individual treatment or as part of a higher order interaction (p>0.05).

64 a) ‘Grass’ seed mix 1.8 1.6 1.4 1.2 1 2009 0.8 2010 0.6 2011 Butterfly species richnessspeciesButterfly 0.4 0.2 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing b) ‘Grass & Legume’ seed mix 3.5

2.5

2 2009 1.5 2010

1 2011 Butterfly species richnessspeciesButterfly 0.5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing c) ‘Grass, Legume & Forb’ seed mix 2.5

1.5 2009 1 2010 2011 Butterfly species richnessspeciesButterfly 0.5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 44. The response of butterfly species richness at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

H6: Establishment of deep rooted species will enhance soil structure and reduce nutrient loss via leaching under either grazing or cutting management systems.

Overall summary: Partially in contradiction to the expectations of hypothesis 6, the more diverse ‘grass, legume & forb’ seed mix was seen to have the highest rates of phosphorus loss within the water leachate. The extent to which phosphorus was leached from the soil for this seed mix increased over time, suggesting this may be an artefact of lower nutrient levels within the soils of simpler seed mixes. While an effect of seed mix was found for the leaching of phosphorus, a similar effect for nitrogen was not seen although again this provided no evidence in support of hypothesis 6. Shifts in the fungal to bacterial ratio (PLFA analyses), which point to an improved capacity of soils to self-regulate their nutrient cycles, were not identified in response to establishing seed mix, whether plants were deep rooted or not. Deep rooting plants, particularly those found within the ‘grass, legume & forb’ seed mix, had no effect on the bulk density of the soil as recorded at 0-10 cm and 10-20 cm horizons. Indeed there was no overall difference in soil bulk density between any of the seed mixtures suggesting that deep rooted plants had no utility as a mitigation measure alleviating soil compaction. There was some indication that the pressure required to penetrate the soil surface may have been slightly higher where more diverse seed mixtures were sown, although this was likely due to a thick mat of roots establishing where the higher overall biomass of plants was greatest within the ‘grass & legumes’ and ‘grass, legume & forb’ plots. The only evidence for a positive effect of establishing seed mix as a method for alleviating soil compaction was found at Jealott’s Hill, where the rate of increase in pressure required to penetrate the soil was lower for the ‘grass & legumes’ and ‘grass, legume & forb’ seed mixes. This at least points to deeper rooting plants in these two seed mixtures having a positive effect on soil structure thus supporting hypothesis 6.

Statistical note for North Wyke: Analysis Part 1: Effects of seed mix and sward management on TON and TP. To test the effects of seed mix on soil structure and nutrient loss via leaching under either grazing or cutting management practises in a fully factorial manner. Analysis was done on the seed mix and management treatments and a sub set of the other treatments; only one level of timing of management (Typical) and cultivation (MC) was available so both of these were excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management and Year. TON and TP data was either log or square root transformed, penetrometer data was log transformed to meet the assumptions of the ANOVA.

Analysis Part 2: Effects of seed mix, cultivation and sward management on TON and TP. To test the effects of seed mix and seed bed preparation on soil structure and nutrient loss via leaching under either grazing or cutting management practises in a fully factorial manner. Analysis was done on the management and cultivation treatments and a sub set of the other treatments; two levels of seed mix (G & GLF) and one level of the timing of management (Typical) were available. Therefore timing of management was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Cultivation and Year. TON and TP data was either log or square root transformed, penetrometer data was log transformed to meet the assumptions of the ANOVA.

Statistical note for Jealott’s Hill: Penetrometer readings, measures of bulk density and all soil chemical properties (e.g. Phosphorus) were recorded in the typically managed experimental plots. ANOVA models tested all interactions of Seed, Management, Cultivation and Year (Timing, i.e. typical vs rested, was redundant for these tests). Samples were recorded in years 2009, 2010 and then again in 2012 at the end of the sampling period.

Soil nutrient loss: total oxidised nitrogen (TON)

Total Oxidized Nitrogen (TON) is the sum of both nitrate and nitrites in the soil. Note that while the 1991 EU Nitrates Directive (91/676/EC) provides a threshold of 50 mg l-1 of nitrates as threshold for polluted groundwater, all concentrations reported at North Wyke site fall considerably below this threshold.

North Wyke: Analysis Part 1) Effects of seed mix and sward management on TON. No treatment effects of management or seed mixture were found on TON concentration in the drainage water during any of the three sampling occasions (November, January and March) in the minimal cultivated plots. Year did have a significant effect for the November (Year: F3,54=15.41, p=<0.001) and January (Year: F3,54=36.55, p=<0.001) leachate samples. Generally TON concentrations in the November and January samples tended to be higher in the third and fourth years of the experiment.

North Wyke: Analysis Part 2) Effects of seed mix, cultivation and sward management on TON. There were no individual cultivation, seed mix or management treatment effects for TON concentrations in the November leachate samples but concentrations were influenced by the interaction between seed mix, management and year and year was significant (Seed mix x Man. x

Year: F3,72=4.05, p=0.034; Seed: F1,3=0.42, p=0.564; Man: F1,6=3.09, p=0.129; year: F3,72=23.71, p=<0.001) and the interaction between seed mix and management (F1,6=10.44, P=0.018). Although not significant, under the cutting management the ‘grass, legume & forb’ sward tended to produce higher TON concentrations in January than the ‘grass only’ sward with the reverse trend found under the grazing management. Within the November TON dataset there is a lot of ‘noise’ but overall concentration levels were lowest in the final year (2012) and highest in 2011 (Fig. 45). Sward management did have a significant effect in the January samples with higher TON concentrations found in the cut plots compared with the grazed plots (Man: F1,6=8.44, p=0.027) but the seed mix and cultivation treatments had no significant effects. No significant treatment effects were found for TON concentrations in the March samples. Year did have a significant effect on TON levels for

January (Year: F3,72=60.28, p=<0.001) and March (Year: F3,72=8.89, p=<0.001). Again there was a considerable amount of ‘noise’ in both the TON January and March datasets, however water samples collected in the first year (2010) and final year (2013) tended to produce lower concentrations than the middle years of the experiment. 1.8 1.6

1.4 1 - 1.2 1 2009 0.8 2010

Nov TON TON mgl Nov 0.6 2011 0.4 0.2 2012 0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 45. Effect of seed mix, cultivation and sward management on TON concentration in leachate samples collected in November at North Wyke.

Soil nutrient loss: total phosphorus (TP)

Total phosphorus (TP) contains three components: soluble reactive phosphorus (SRP), soluble unreactive or soluble organic phosphorus (SUP) and particulate phosphorus (PP). Unlike nitrates there is currently no EU legislative threshold for TP. However, under the European Water Framework Directive (Directive 2008/105/EC), watercourses are classified as having ‘good’ water quality if they contain between 0.04 and 0.12 mg SRP L-1 (depending on the alkalinity and altitude of the watercourse) and the Environment Agency (EA) class rivers with phosphate levels greater than 0.1 mg SRP L-1 as having high concentrations (http://www.environment- agency.gov.uk/research/planning/34383.aspx). Note, the majority of TP concentrations reported for the North Wyke site fall below the EU and EA SRP values considered to be of high concentrations. Typically TP concentrations at North Wyke were below 40 µg l-1 (0.04 mg l-1) although there were a few outliers (appendix 1c). North Wyke: Analysis part 1) Effects of seed mix and sward management on TP. Management had a significant effect on TP concentration in the November samples (Man: F1,9=12.51,P=0.006) with higher concentrations generally found under the grazing regime particularly in the GL and GLF swards (Fig. 46). Although the seed mix treatment was not found to be significant, overall highest losses of TP were from the GLF swards. TP concentrations in November tended to increase in the later stages of the experiment (Year: F3,54=18.04, p=<0.001). 400 350

300 1 - 250 200 2009

150 2010 Nov TP µg µg TP Nov l 100 2011 50 2012 0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 46. Effect of seed mix and sward management on TP concentration in leachate samples collected in November at North Wyke.

There were significant interactions between seed mix and year (Seed x Year: F6,54=2.81, p=0.036; Seed: F2,6=0.11, p=0.897; year: F3,54=29.42, p=<0.001) on TP concentration in the leachate samples collected in January (Fig. 47). The lowest levels of TP tended to be in the first samples collected in January 2010. Overall the highest January TP mean concentrations were produced by the GL sward

68 in 2010 and 2011, the GLF sward in 2012 and G sward in 2013. Management had no significant effect on TP content in the January samples.

140 120

100

1 - 80 2010 60

Jan TP TP µlJan 2011 40 2012 20 2013 0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 47. Effect of seed mix and sward management on TP concentration in leachate samples collected in January at North Wyke.

There were significant interactions between management and year (Man. x Year: F3,54=3.83, p=0.034; Man: F1,9=8.93, p=0.015; Year: F3,54=145.47, p=<0.001) on TP concentration in the leachate samples collected in March (Fig. 48). The lowest levels of TP were found in the first samples collected in March 2010. Generally the grazing management produced higher March TP levels in 2010 and 2011 than the cutting management with the reverse trend found in 2012 and 2013. Seed mix had no significant effect on TP content in the March samples. 250

200

1 - 150 2010 100

2011 Mar TP TP l µg Mar 50 2012 2013 0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 48. Effect of seed mix and sward management on TP concentration in leachate samples collected in March at North Wyke.

North Wyke: Analysis part 2) Effects of seed mix, sward management and cultivation on TP. The minimum cultivation plots produced higher TP concentrations in the November leachate water than the ploughed plots (Cult: F1,12=7.56, p=0.018). Although seed mix did not have a significant effect overall the GLF plots produced higher TP concentrations in November than the G only plots. TP

November concentrations were also affected by year (Year: F3,72=19.62, p=<0.001) with levels tending to increase throughout the duration of the experiment (Fig. 49). Management had no significant effect on TP content in the November samples 400 350

300 1 - 250 200 2009

150 2010 Nov TP ug l ug TP Nov 100 2011 50 2012 0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 49. Effect of seed mix, cultivation and sward management on TP concentration in leachate samples collected in November at North Wyke.

There were no individual cultivation, seed mix or management treatment effects for TP concentrations in the January leachate samples but concentrations were influenced by the interaction between seed mix, cultivation and year (seed mix x Cult. x year: F3,72=3.62, p=0.036; seed mix: F1,3=0.35, p=0.595; Cult: F1,12=3.63, p=0.081; year: F3,72=47.20, p=<0.001). Although not significant, overall the ‘grass, legume & forb’ sward produced higher January TP concentrations than the ‘grass only’ sward in both seed bed preparation methods. Lowest TP concentrations were found in the establishment year (2009) and highest in the final year (2012)(Fig. 50). 140 120

100

1 - 80 2010 60

Jan TP TP Jan ug l 2011 40 2012 20 2013 0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 50. Effect of seed mix, cultivation and sward management on TP concentration in leachate samples collected in January at North Wyke

As with the November and January TP concentrations, the plots sown with the ‘grass, legume & forb’ seed mix produced higher TP concentrations in the March leachate water than the ‘grass only’ plots although this effect was not significant. TP March concentrations were also affected by year (Year:

F3,72=38.57, p=<0.001) with levels generally increasing in the later stages of the experiment (Fig. 51). The minimally cultivated plots produced higher TP concentrations compared with the ploughed plots

(Cult: F1,12=5.72, p=0.034). There were no significant management effects. 250

200

1 - 150 2010 100

2011 Mar TP TP l ug Mar

50 2012 2013 0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 51. Effect of seed mix, cultivation and sward management on TP concentration in leachate samples collected in March at North Wyke

Jealott’s Hill: See above comment.

Soil structure

Bulk density (0-10cm)

North Wyke: Analysis part 1) Effects of seed mix and sward management on bulk density (0- 10cm). No significant response to seed mix or subsequent management was found on soil bulk densities in the 0-10cm horizon. Year did have a significant effect (Year: F2,37.5=5.39, p=0.009) with the highest 0-10cm bulk densities tending to occur in 2010 (Fig. 52).

0.90

) 3 - 0.80 0.70 0.60 0.50

0.40 2009 10cm horizoncm (g

- 0.30 2010 0.20 2012 0.10 0.00 MC MC MC MC MC MC

Bulk density0 in Bulk G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 52. Effect of seed mix and sward management on soil bulk density (0-10cm horizon) at North Wyke.

North Wyke: Analysis part 2) Effects of seed mix, sward management and cultivation on bulk density (0-10cm). Soil bulk density was significantly lower in the 0-10cm horizon where minimal cultivation rather than ploughing was used to establish the seed mixes (Cult: F1,33.9=29.13 p=<0.001) (Fig. 53). Year had a significant effect with soil bulk density tending to be lowest in the establishment year (2009) and highest in 2010 (Year: F2,48.2=7.38, p=0.002). Seed mix and the management treatment had no significant effect on soil bulk density in the 0-10cm horizon.

1.00

) 3 - 0.90 0.80 0.70 0.60 0.50 2009

10cm horizon (g cm (g horizon10cm 0.40 - 0.30 2010 0.20 2012 0.10 0.00

Bulk density in 0 in density Bulk MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 53. Effect of seed mix, cultivation and sward management on soil bulk density (0-10cm horizon) at North Wyke.

Jealott’s Hill (Bulk density 0-10 cm): The bulk density of soil from the 0-10 cm horizon of soil respond to neither seed mix, management, cultivation or year (p>0.05).

Bulk density (10-20cm)

North Wyke: Analysis part 1) Effects of seed mix and sward management on bulk density (10- 20cm). Bulk density in the 10-20cm horizon was significantly higher where swards were cut rather than grazed (Man. F1,27.9=6.95, p=0.014). Year had a significant effect with soil bulk density increasing throughout the duration of the experiment (Year: F2,38.4=10.46, p=<0.001)(Fig. 54). Seed mix had no significant effect on bulk density in the 10-20cm horizon.

1.20

)

3 - 1.00

0.80

0.60

2009 20cmcm (g horizon - 0.40 2010

0.20 2012

0.00 MC MC MC MC MC MC

Bulk density in 10 in density Bulk G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 54. Effect of seed mix and sward management on soil bulk density (10-20cm horizon) at North Wyke.

North Wyke: Analysis part 2) Effects of seed mix, sward management and cultivation on bulk density (10-20cm). In contrast to the 0-10cm horizon, bulk density was significantly lower in the 10-

20cm horizon where ploughing was used to establish the seed mixes (Cult: F1,33.4=36.76, p=<0.001) (Fig. 55). Bulk densities were significantly higher under the cutting management compared with the grazing regime (Man: F1,33.4=8.77, p=0.006). Year had a significant effect with soil bulk density generally increasing throughout the duration of the experiment (Year: F2,50.3=11.28, p=<0.001). Seed mix had no significant effect on soil bulk density in the 10-20cm horizon.

1.20

)

3 - 1.00

0.80

0.60

2009

20cm horizoncm (g - 0.40 2010

0.20 2012

0.00

Bulk density10 in Bulk MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 55. Effect of seed mix, cultivation and sward management on soil bulk density (10-20cm horizon) at North Wyke.

Jealott’s Hill (Bulk density 10-20 cm): The bulk density of soil from the 10-20 cm horizon of soil did not respond to either seed mix, management or cultivation (p>0.05), although there was a significant difference between years (F2,141=60.9, p<0.001). There was some evidence that bulk

73 density was slightly lower for this soil profile by 2012 (mean=1.21, ±SE=0.03) relative to that seen in 2010 (mean=1.45, ±SE=0.02). Note that at Jealott’s Hill bulk density was assessed at this deeper profile only in 2010 and 2012. This year effect is in agreement to some extent with the penetrometer data presented above which suggested an overall reduction in soil compaction to a depth of 49 cm over time.

Soil penetrometer resistance

North Wyke: Analysis part 1) Effects of seed mix and sward management on soil penetrometer resistance: Within the minimum cultivation plots no significant response to seed mix was found in penetrometer resistance. Year had a significant effect (Table 3) throughout the soil profile where penetrometer resistance was highest in 2010 and lowest in 2012, the exception being at a depth of 3.5cm where resistance was highest in 2012 and lowest in 2010. The summer of 2012 was wetter than the previous years, therefore lower resistance in 2012 is likely to be a consequence of higher soil moisture content than in 2009 and 2010. At many depths there was a significant interaction between management and year although the management treatment itself was only found to be significant at depths of 7cm and 28cm where resistance was generally highest under the grazing regime. The management and year interaction showed that while in 2009 and 2010 the amount of pressure required to penetrate the soil at all depths was greatest in the grazed plots, by the final year resistance was higher in the cut plots (Fig. 56).

Depth (cm) Factor Probability

3.5 year F2,34.1=9.49, p=<0.001 7 year F2,54=5.64, p=0.006 management F1,54=8.87, p=0.004 year x management F2,54=8.24, p=<0.001 10.5 year F2,54=34.53, p=<0.001 year x management F2,54=8.83, p=<0.001 14 year F2,54=54.29, p=<0.001 year x management F2,54=10.29, p=<0.001 17.5 year F2,54=60.38, p=<0.001 year x management F2,54=42.08, p=<0.001 21 year F2,54=60.38, p=<0.001 year x management F2,54=7.42, p=<0.001 24.5 year F2,54=68.11, p=<0.001 year x management F2,54=12.2, p=<0.001 28 year F2,54=76.35, p=<0.001 management F1,54=6.41, p=0.014 year x management F2,54=8.83, p=<0.001 31.5 year F2,37=87.72, p=<0.001 year x management F2,37=7.36, p=0.002 35 year F2,38.2=70.76, p=<0.001 38.5 year F2,38.6=66.49, p=<0.001 42 year F2,54=73.19, p=<0.001 45.5 year F2,54=92.12, p=<0.001 49 year F2,38.3=109.29, p=<0.001 year x management F2,38.3=3.47, p=0.041

Table 3. Summary of significant results of seed mix and sward management effects on soil penetrometer resistance in minimally cultivated plots at North Wyke

2009 2010 mean pressure (bar) mean pressure (bar) 0 10 20 30 40 0 20 40 60 0 0

10 10

20 20

Depth Depth 30 30 (cm) (cm)

40 40

50 50

60 60

2012 mean pressure (bar) 0 10 20 30 0

20 MC cut Typical MC grazed Typical Depth (cm) 30

Fig. 56. Effect of sward management on soil penetrometer resistance at North Wyke.

North Wyke: Analysis part 2) Effects of seed mix, cultivation and sward management on soil penetrometer resistance. The grass only and grass, legume forb seed mixtures did not have a significant effect on soil resistance. Year had a significant effect (Table 4) throughout the soil profile with penetrometer resistance highest in 2010 and lowest in 2012, the exception being at a depth of 3.5cm where resistance was highest in 2012 and lowest in 2010. The summer of 2012 was wetter than the previous years, therefore lower resistance in 2012 is likely to be a consequence of higher soil moisture content than in 2009 and 2010. The management treatment had a significant effect on soil resistance at depths of 7cm, 10.5cm, 14cm and 24.5cm with the grazed plots having higher resistance values than the cut plots. At many depths there was a significant interaction between management and year (Table 4) with higher soil resistance measurements in the grazed plots in 2009 and 2010 compared with the cut plots with the reverse trend in 2012. Significant cultivation effects were found at depths of 7cm, 17.5cm, 21cm and 24.5 cm with generally greater soil resistance in the minimum cultivation plots than the ploughed except at the 7cm depth. In 2009 and 2010 soil resistance was lower in the 14-24.5cm layer below ground in the ploughed treatment compared with the minimally cultivated plots, as would be expected, however the upper soil layer (7-17.5cm) showed a greater resistance in the ploughed plots in the final year of the experiment (Fig. 57).

Depth (cm) Factor Probability

3.5 year F2,47.2=7.73, p=<0.001 7 year F2,72=9.94, p=<0.001 cultivation F1,72=10.12, p=0.002 management F1,72=10.32, p=0.002 year x management F2,72=14.93, p=<0.001 10.5 year F2,72=24.74, p=<0.001 management F1,72=7, p=0.01 year x cultivation F2,72=6.3, p=0.003 year x management F2,72=12.47, p=<0.001 14 year F2,72=39.37, p=<0.001 management F1,72=2.95, p=0.09 year x cultivation F2,72=7.25, p=0.001 year x management F2,72=14.19, p=<0.001 17.5 year F2,72=39.71, p=<0.001 cultivation F1,72=27.67, p=<0.001 year x cultivation F2,72=5.94, p=0.004 year x management F2,72=9.52, p=<0.001 21 year F2,72=40.78, p=<0.001 cultivation F1,72=25.4, p=<0.001 year x management F2,72=7.71, p=<0.001 24.5 year F2,72=58.52, p=<0.001 cultivation F1,72=6.53, p=0.013 management F1,72=4.45, p=0.038 year x management F2,72=6.25, p=0.003 28 year F2,72=130.61, p=<0.001 year x management F2,72=17.2, p=<0.001 31.5 year F2,50.1=111.75, p=<0.001 year x management F2,50.1=10.42, p=<0.001 35 year F2,50.4=76.59, p=<0.001 year x management F2,50.4=4.72, p=0.013 38.5 year F2,50.6=76.43, p=<0.001 42 year F2,50.9=98.05, p=<0.001 year x management F2,50.9=4.39, p=0.017 45.5 year F2,51.7=94.24, p=<0.001 49 year F2,51.3=130.54, p=<0.001 year x management F2,51.3=4.93, p=0.011

Table 4. Summary of significant results of seed mix, cultivation and sward management effects on soil penetrometer resistance at North Wyke

2009 2010 mean pressure (bar) mean pressure (bar) 0 20 40 0 20 40 60 0 0

10 10

20 20

Depth Depth 30 30 (cm) (cm)

40 40

50 50

60 60

2012 mean pressure (bar) 0 10 20 30 0

20 MC Typical Ploughed Typical Depth (cm) 30

Fig. 57. Effect of cultivation on soil penetrometer resistance at North Wyke.

Jealott’s Hill (Penetrometer): The penetrometer study involved recording the pressure (kg cm-2) required to penetrate the soil using a 70cm steel probe to 3.5, 7.0, 10.5, 14.0, 17.5, 21.0, 24.5, 28.0, 31.5, 35.0, 38.5, 42.0, 45.5, 49.0 cm depths at 14 locations per plot. A logarithmic curve (y = α +

β.loge(x)) was fitted to the penetrometer data. The intercept (α) is the pressure required to penetrate the soil surface (0-3.5 cm soil depth). The slope coefficient, (β), is the rate with which

78 pressure required to penetrate the soil increases with depth. As the value of β increases, i.e. the slope of the curve becomes steeper, the rate with soil resistance to penetration increases with depth becomes greater (upper right hand curve in Figure), or vice versa shown by the (lower right hand curve). The logarithmic function was a good fit to all data at both Jealott’s Hill, with R2 values typically in excess of 0.80. The response of α and β to the fixed effects of Seed, Management, Cultivation and Year were tested using repeated measures ANOVA.

β increases: soil 80 compaction increases 70 y = 22.3ln(x) + 7.01 with depth at a greater 60 rate 50 80 40 70 > β 30 60 y = 11.1ln(x) + 3.65 20 50 Pressure 10 40 (kg cm-2) 0 30 0 5 10 15 20 10 80 0 < β 70 β decreases: 0 5 10 15 60 50 indicating reduced y = 5.98ln(x) + 5.61 Soil depth interval 40 rate of soil 30 compaction 20 increase with depth 10 0 0 5 10 15

Logarithmic curves (y = β.ln(x) + α) describe rates of change in soil penetrability. The slope β is derived for each experimental plot and then tested against the fixed effects of Seed mix, management, cultivation and year.

Jealott’s Hill Soil penetrability at soil surface: Penetrometer intercept (α): Soil compaction at the surface (0-3.5 cm depth) for the Jealott’s Hill site (as defined by the intercept (α) of a logarithmic function fitted to the penetrometer data) showed a significant response to the interaction between establishing seed mixture and the number of years since establishment (seed × year: F2,124=16.6, p<0.001; seed: F2,6=10.8, p<0.01; year: F1,124=11.1, p<0.01). The amount of pressure required to penetrate this first 3.5 cm of the soil was initially similar for all three seed mixtures in 2009, the first year following grassland establishment in the autumn of 2008. However, soil compaction for the two most complex seed mixtures (‘grass & legumes’ and ‘grass, legume & forb’) increased above that seen for the ‘grass’ seed mixture in 2010, two full years into the establishment of the swards (Fig. 58). This increase in apparent pressure required to penetrate the soil surface in 2010 may be a product of the development of a dense surface mat of legume roots in these inherently higher productivity swards. However, by the final year of the study (2012) this increase in soil compaction linked with the inclusion of legumes in the seed mix had largely disappeared, with all three seed mixes requiring approximately similar levels of pressure to penetrate the soil surface. Again, this links to the general collapse in cover of legumes across all plots. For the ‘grass & legumes’ and ‘grass, legume & forb’ seed mixes this pressure had fallen noticeably between 2010 and 2012, although from 2009 to 2012 the pressure showed no change for the ‘grass’ seed mix. It is likely that this collapse in pressure required to penetrate the soil surface in the ‘grass & legumes’ and ‘grass, legume & forb’ seed mixes is linked to at least a general reduction in legume cover (see below) over the four full years of sward establishment. No other significant effects on surface soil compaction were identified, the most surprising of which is an absence of an effect of soil cultivation, either individually or as part of a significant interaction with either seed mix or management (p>0.05).

intercept) α α 8 2009 6 2010 4 2012

2 Surface compaction ( compaction Surface 0 Grass GL GLF Seed mix

Fig. 58. Effect of establishing seed mix on surface soil compaction (the pressure required to penetrate the first 3.5 cm of the soil surface) at Jealott’s Hill, as represented by α of the logarithmic curve fitted to soil penetrometer data.

Jealott’s Hill Rate of increase in soil compaction with depth (β): Overall soil compaction was described by the slope parameter β of a logarithmic function fitted to the penetrometer data. The β slope defines the rate of increase in pressure required to penetrate the soil with depth to 49.0 cm from the surface. This index of soil compaction was found to differ between the establishing seed mixes (seed: F2,6=5.98, p=0.03; Fig 59), so that the inclusion of legumes in seed mixtures had a positive effect on soil structure. Although establishing swards with ‘grass & legume’ and ‘grass, legume & forb’ seed mixes had an overall positive effect on soil compaction, there was no evidence that these two seed mixes differed between each other. It was predicted that deeper rooting plants in the ‘grass, legume & forb’ seed mix would promote an overall greater reduction in soil compaction, however this does not seem to be the case. While it is possible that the structural properties of the clay soil at this site may require longer periods before amelioration by sown plants can have an effect (Raty et al., 2010) the absence of a trend after 4 years would point to the limited utility of the forb component in the seed mix as a mitigation measure against soil compaction. In addition to the effect of establishing seed mix, soil cultivation used in initial seed bed preparation in 2008 also affected the rate of increase in soil compaction with soil depth as part of an interaction with sward management and year (cult. × man. × year: F6,86=430, p<0.001; cult. F1,22=5.51, p=0.03; man. F1,11=0.19, p>0.05; year: year F2,86=102.6, p<0.001; Fig. 60). Across all combinations of sward management and seed bed cultivation there is a clear reduction in overall soil compaction (lower β slope parameter) between 2009 and 2010, with this loss in soil compaction being lost to some extent by 2012 although still remaining lower that was seen in 2009. It should be noted, however, that differences in soil water content between years may have a key factor that affected the penetrability of the soil. Year has been retained in models, although the interpretation of between year differences should be treated with caution due to lack of control over soil moisture content. However, after four years of sward establishment (2012) differences between treatments suggests that the deep ploughing cultivation methods used to establish seed beds, either where cutting or grazing management was applied to the sward, would result in increased levels of soil compaction relative to the shallow cultivation methods. This longer term negative impact of deep cultivation methods reflects previous findings that suggest that mechanically loosened soils are prone to recompaction (Munkholm et al., 2005).

4 slope)

β β 3

2 depth ( depth 1

Rate of increase in soil ofsoil increase Ratein compaction with Grass GL GLF Seed mix

Fig. 59. Differences in soil compaction in response to establishing seed mix. Soil compaction is defined by the rate of increase in pressure required to penetrate the soil to a depth of 49.0 cm (β slope parameter of the logarithmic function fitted to the penetrometer data).

10 β) 8

6 2009 4 2010

2 2012 depth (Slope parameter (Slope depth 0

Deep Shallow Deep Shallow Rate of increase in soil compaction with compaction soil in increase of Rate Cut Cut Graze Graze Management and cultivation of seed bed

Fig. 60. Differences in soil compaction in response to the interaction between sward management practice, the cultivation method used in initial seed bed preparation in 2008 and the subsequent changes in these factors from 2009 to 2012. Soil compaction is defined by the rate of increase in pressure required to penetrate the soil to a depth of 49.0 cm (β slope parameter of the logarithmic function fitted to the penetrometer data). For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

PLFA Fungal to bacterial ratio.

Jealott’s Hill: PLFA fungal to bacterial ratios were only assessed in 2012, the final sampling year of the study. This was intended to give the soils enough time to establish differences in response to sward management and establishing seed mixture. There were however no significant effects, either individually or as part of interaction terms between establishing seed mix, sward management and cultivation technique. This suggested that there has been no shift towards increased self-

81 regulation of soil nutrient cycles within these grasslands (Bardgett & McAlister 1999; De Vries et al 2012). The absence of a management effect is in agreement with previous studies that have failed to identify a role in the cessation of cutting or grazing on fungal: bacterial ratios (Bardgett & McAlister 1999). However, as plant functional traits linked with fast growing and nitrogen exploitative survival strategies have been shown to select for bacterial dominated communities (Orwin et al 2010), the shift in floral composition away from those typical of improved grasslands characterised by ‘grass & legume’ and ‘Grass, Legume & Forb’ seed mixtures was expected to increase fungal dominance. The failure over a five year period to see such shift suggests that reductions in soil nitrogen losses promoted by high fungal: bacterial ratio would be unlikely to occur for these swards (De Vries et al 2012). This has implications for both sustainable food production as well as the capacity of these grasslands to mitigate against diffuse pollution (De Vries et al 2012).

H7: Legume or legume plus forb rich swards will increase grassland productivity and forage quality compared with very low input grass dominated swards.

Overall summary: This hypothesis was accepted, with higher yields of silage and increased nutrient quality of the herbage being found where legumes and non-legume forbs were part of the establishing seed mixture. Although the inclusion of the non-legume forb component resulted in slightly higher herbage production and quality over simply the presence of legumes, this effect was comparatively small when compared to the benefits of including legumes over a sward sown with just grasses. While the inclusion of legumes and non- legume forbs did not increase the daily rate of increase in livestock weight, they did result in swards that could be grazed for longer periods of time.

Statistical note for North Wyke: To investigate the effects of adding legumes and non-legume forbs to the sward on forage yield and quality, analysis was carried out on the seed mix used for the period 2010 – 2012 based on two cuts: three levels of seed mix (G, GL and GLF) were available in the minimal cultivation plots and two levels of seed mix (G and GLF) in the ploughed plots. Forage yields were only assessed on the cut plots. All analyses were carried out on the typically managed plots – this model therefore identifies significant effects of all possible interactions between Seed mix, Management and Year. Animal performance was only assessed on the grazed plots. All analyses were carried out on the available unit area, with animals excluded for a short rest period in the summer, and allowed to graze this area at the end of the exclusion period. This model therefore identifies significant effects of Seed mix for each year.

Statistical note for Jealott’s Hill: Note that as all herbage analyses originates from data derived from experimental plots managed by cutting and not grazing, ‘management’ as a treatment is not relevant in these analyses and has been ignored. For comparability all chemical analysis of the Herbage are derived from the first sward cut in the year as this was common to both typical and rested timings of management, with the second sward cut being absent under rested management. However, in the case of dry matter yield (tonnes ha- 1) this was represents the summed mass of silage removed from an experimental plot over the course of a year, and so in the case of typical management timings this is the combination of both the early and late sward cut.

Dry matter yield from minimal cultivation cut plots

North Wyke: No significant effect of seed mix used was found on the production of total dry matter for the period 2010-2012 in the minimal cultivation plots (F2,6 = 0.23, p = 0.80) (Fig. 62) (G 7.56±0.38 t

-1 -1 -1 ha . GL 7.94±0.27 t ha , GLF 7.43±0.57 t ha ), with no significant differences at the first cut (F2,6 = 0.03, p = 0.969) or second cut (F2,6 = 1.37 P = 0.324).

Dry matter yield from ploughed cut plots

North Wyke: No significant effect of seed mix used was found on the production of total dry matter -1 for the period 2010-2012 in the ploughed plots (F1,3 = 4.14, p = 0.135 G 6.05±0.3 t ha , GLF 7.6 ±0.5 t ha-1 (Fig. 61).

Fig. 61. Total dry matter yield production (t ha-1) from two cuts combined (mean ± SE mean).

Jealott’s Hill (Dry matter yield from cut plots): The total dry matter yield obtained from cutting management at Jealott’s Hill was directly affected by an interaction between the establishing seed mix and the number of years of plot establishment (seed × year: F6,135=7.1, p<0.001; seed: F2,6=62.7, p<0.001; year: F3,135=73.8, p<0.001; Fig. 62). In the initial establishment year of 2009 the dry matter yield was consistently higher where legumes (‘grass & legume’ and ‘grass, legume & forb’) were included in the seed mixes in addition to just grass. Indeed, for 2009 the yield could be as much 5 tonnes ha-1 greater than the 3 tonnes ha-1 seen for the ‘grass’ only seed mix plots. The general trend across all seed mixes was for the yield to decrease from 2009 to 2012, although the massive decline in cut yield in 2010 is linked with a drought year, rather than being associated with an underlying characteristics resulting from the seed mixes. While the decline in dry matter yield was negligible for the already low yield of the ‘grass’ only seed mix, it was far more pronounced where legumes were part of the seed mix dropping from c. 8 tonnes ha-1 in 2009 to 3 tonnes ha-1 by 2012. This collapse in yield was most evident (with the exception of the extreme drought year in 2010) between 2011 and 2012 and is likely the direct result of the wide scale collapse in populations of the sown component of legumes by the fourth year of plot establishment.

) 1 - 10

8 2009 6 2010 4 2011 2012

2 Dry matter yield (tonnes ha (tonnes yield matter Dry

0 Grass GL GLF Seed mix

Fig. 62. Effect of establishing seed mix and its interaction with year on the summed yearly dry matter yield of the cut hay for Jealott’s Hill.

In addition to the overall effect of the interaction between establishing seed mix and year, the dry matter yield was also affected by the timing of cutting management operations, effectively whether there was one or two cuts per year (timing: F1,11=7.72, p=0.02; Fig. 63). As may be expected the addition of a second cut contributing positively to the total dry matter yield resulting in an overall increase of c. 0.5 tonnes ha-1. The relatively small overall size of this difference in yield reflects the fact that it is averaged over the four year monitoring period and across all seed mixes, and so hides the differences described above for the year and seed mix interaction. Shallow cultivation of the seed bed in 2008 also had a small overall benefit for dry matter yield, increasing it by c. 0.5 tonnes -1 ha over the four year period (cultivation: F1,23=7.18, p=0.01; Fig. 64).

) 1 - 5

1 Dry matter yield (tonnes ha (tonnes yield matter Dry

0 Typical Rested Timing of management

Fig. 63. Effect of timing of cutting management operation on the summed yearly dry matter yield of the cut hay for Jealott’s Hill.

) 1 - 5

1 Dry matter yield (tonnes ha (tonnes yield matter Dry

0 Deep Shallow Cultivation

Fig. 64. Effect of initial depth of seed bed cultivation in 2008 on the summed yearly dry matter yield of the cut hay for Jealott’s Hill. For seed bed cultivation , deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Forage quality

North Wyke: Forage quality from minimal cultivation plots. No significant effect of seed mix was found on nitrogen content at the first sampling in June (F2,6=0.96, p = 0.434), whilst a significant -1 difference was recorded at the second sampling in August (F2,6=6.42, p=0.032, G 18.8±1.05 mg kg , GL 21.11±1.07 mg kg-1, GLF 21.14±1.04 mg kg-1). Total herbage N content was significantly higher in the grazed plots compared with the cut plots in all years in both the first silage cut in June (Year x

Man: F2,36=15.37, p = <0.001; Year: F2,36=7.76, p = 0.005; Man: : F1,9=133.75, p=<0.001) and the second silage cut in August (Year x Man: F2,36=9.14, p = 0.002; Year: F2,36=45.70, p = <0.001; Man: F1,9=65.33, p=<0.001). No significant effect of seed mix was recorded on phosphorus content at either sampling (first sampling F2,6=0.33, p=0.730; second sampling F2,36=1.78, p=0.248). The phosphorus content of the herbage was significantly affected by the sward management in both

June (Man: F1,9=89.66, p=<0.001) and August (Man: F1,9=21.76, p=0.001) with higher concentrations recorded in the grazed plots. No significant effect of seed mix was found on calcium content at the -1 first sampling (F2,6=1.34, p=0.330) or the second sampling (F2,6=4.70, p=0.059 G 5.93±0.16 mg kg , GL 7.34±0.51 mg kg-1, GLF 7.00 ±0.36 mg kg-1). No significant effect of seed mix was recorded on magnesium content at the first sampling (F2,6=0.47, p=0.644) or second sampling (F2,6=3.04, p=0.059), or on sodium content at the first sampling (F2,6=1.47, p=0.302) or second sampling (F2,6=4.77, p=0.058). Herbage magnesium content was significantly higher in the grazed plots than the cut plots in June (Man: F1,9=16.23, p=0.003) but not in August. Higher levels of herbage sodium content were recorded in the cut plots in both (Man: F1,9=5.54, p=0.043) and August (Man: F1,9=8.35, p=0.018). Pepsin cellulose digestibility (DOMD) was not significantly affected by seed mix at the first -1 sampling (F2,6=2.36, p=0.176), but was at the second sampling (F2,6=6.03, p=0.037, G 596.2±5.2 g kg , GL 619.3±6.5 g kg-1, GLF 618.0±5.9 g kg-1). DOMD herbage content was significantly higher in the grazed plots than the cut plots in June (Man: F1,9=15.16, p=0.004) however sward management had no significant effect in August on DOMD.

North Wyke: Forage quality from ploughed plots. A significant effect of seed mix was recorded in -1 the ploughed plots on nitrogen at the first sampling (F1,3=89.22, p=0.003 G 13.9±0.8 mg kg , GLF

-1 -1 16.8±0.9 mg kg ), and at the second sampling (F1,3=40.17, p=0.008 G 18.81±0.9 mg kg , GLF 23.4±1.1 mg kg-1). Total herbage N content was significantly higher in the grazed plots compared with the cut plots in all years at the first sampling in June (Year x Man: F2,24=33.34, p =<0.001; Year: F2,24=17.59, p = <0.001; Man: : F1,6=297.40, p=<0.001) and the second silage cut in August (Man: F1,6=115.97, p=<0.001). A significant effect of seed mix was recoded on phosphorus content at the first sampling -1 -1 (F1,3=11.16, p=0.044 G 2.02±0.12 µg kg , GLF 2.18±0.10 µg kg ), but not at the second sampling (F1,3=0.23, p=0.66). The phosphorus content of the herbage was significantly affected by the sward management in both June (Man: F1,6=181.50, p=<0.001) and August (Man: F1,6=27.44, p=0.002) with higher concentrations recorded in the grazed plots. Herbage calcium content was significantly higher in the more diverse swards (GLF) than the ‘grass only’ swards throughout the duration of the experiment in both June and August (June; Year x Seed: F2,24=10.44, p =<0.001; Year: F2,24=18.03, p = <0.001; Seed: F1,3=30.10, p=0.012 and August; Year x Seed: F2,24=11.56, p =0.001; Year: F2,24=33.85, p = <0.001; Seed: F1,3=578.93, p=<0.001) with Ca content declining over time. The calcium content in the August herbage samples was significantly higher in the cut plots than the grazed plots in all years

(Year x Man: F2,24=5.67, p =0.019; Man: F1,6=8.25, p = 0.028) but no such effect was found in the June samples. A significant seed mix and year interaction was recorded on magnesium content at the first sampling (Year x Man: F2,24=8.82, p=0.004; Year: F2,24=22.15, p = <0.001; Seed: : F1,3=32.59, p=0.011) with a pronounced decline over time occurring in the GLF plots (2010 1.57±0.04 mg kg-1, 2011 1.36±0.05 mg kg-1, 2012 1.08±0.09 mg kg-1) but to a lesser extent in the G plots (2010 1.00±0.07 mg kg-1, 2011 1.10±0.03 mg kg-1, 2012 0.89±0.10 mg kg-1). Magnesium content was also significantly affected by seed mix and year at the second sampling (Year x Seed: F2,24=7.49, p=0.008; Year: F2,24=31.04, p =<0.001; Seed: : F1,3=242.80, p=<0.001) with a decline occurring over time being more pronounced in the GFL plots (2010 2.31±0.14 mg kg-1, 2011 2.04±0.15 mg kg-1, 2012 1.41±0.08 mg kg -1) than the G plots (2010 1.40±0.04 mg kg-1, 2011 1.41±0.05 mg kg-1, 2012 1.11±0.05 mg kg-1). Magnesium content in the June herbage samples was significantly higher in the grazed plots than the cut plots in all years (Year x Man: F2,24=5.79, p=0.017; Year: F2,24=22.15, p =<0.001; Man: F1,6=8.13, p=0.029) with the reverse trend found in the August samples (Year x Man: F2,24=4.77, p=0.031; Year: F2,24=31.04, p =<0.001; Man: F1,6=7.19, p=0.036). Sodium content in the June herbage samples was significantly affected by an interaction between year and seed mix (Year x Seed:

F2,24=6.11, p=0.015; Year: F2,24=36.21, p =<0.001; Seed: F1,3=8.66, p=0.060) with higher sodium concentrations found in the GLF seed mix than the G only plots in all years with an increase followed by a decline in both the G plots (2010 0.75±0.10 mg kg-1, 2011 1.45±0.09 mg kg-1, 2012 0.80±0.04 mg kg-1) and GLF plots (2010 1.82±0.27 mg kg-1, 2011 2.33±0.29 mg kg-1, 2012 1.13±0.16 mg kg-1). A significant effect of seed mix on sodium was also found at the second sampling (F1,3=14.35, p0.32 G 0.93±0.06 mg kg-1, GLF 2.07±0.18 mg kg-1). Sodium content was significantly higher in the cut plots compared to the grazed plots in both June (Man: F1,6=14.54, p=0.009) and August herbage samples (Man: F1,6=7.19, p=0.036).

Pepsin cellulose digestibility was significantly affected by seed mix at the first sampling (F1,3=14.05, p=0.033) but not in the second sampling(F1,3=0.49, p=0.534). Sward management also significantly affected digestibility but only in the June herbage samples (F1,6=39.35, p=<0.001) with higher DOMD found in the grazed plots compared with the cut plots.

Jealott’s Hill (Forage quality):

Total herbage nitrogen (N) concentration: Total herbage Nitrogen (% w/w) responded to an interaction between the establishing seed mix and the number of years of plot establishment (seed × year: F6,135=43.6, p<0.001; seed: F2,6=36.6, p<0.001; year: F3,135=190.4, p<0.001; Fig. 65). Total herbage Nitrogen was principally linked to the inclusion of the Nitrogen fixing legumes within the seed mixes, with its percentage in the sward being on average c. 70-80 % higher than that found in

86 the ‘grass’ only plots during 2009. While the ‘grass’ only seed mix maintained the percentage of nitrogen for the four year period at c. 1 % w/w, in the case of both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes the percentage nitrogen dropped by around half from c. 2 to 1 % w/w between 2009-2012. While there no apparent difference in sward nitrogen resulting from the inclusion on non-legume forbs, the decline in legume cover over the four years was likely directly responsible for the collapse in herbage nitrogen.

2.5

1.5 2009 2010 1 2011

Herbage Nitrogen (%) Nitrogen Herbage 0.5 2012

0 Grass GL GLF Seed mix

Fig. 65. Effect of establishing seed mix and its interaction with year on the percentage of Nitrogen in the cut hay for Jealott’s Hill.

There was also a significant effect of the depth of seed bed cultivation used in 2008 on total herbage nitrogen (F1,35=4.47, p=0.04; Fig. 66). The effect of seed bed cultivation on herbage nitrogen was limited with a c. 0.7% w/w increase relative to where deep ploughing has been used. This increase in herbage nitrogen may be linked with the increased retention of total soil nitrogen under the shallow cultivation approach as described above. It should be noted that the relatively small difference in herbage nitrogen and the relatively high level of variation linked with its means indicates that as in importance in terms of applied management is limited.

1.65

1.6

1.55

1.5

1.45 Herbage Nitrogen (%) Nitrogen Herbage 1.4

1.35 Deep Shallow Cultivation

Fig. 66. Effect of cultivation method used during seed bed preparation in 2008 on the percentage of Nitrogen in the cut hay for Jealott’s Hill. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Herbage phosphorus (P) content: The phosphorus content of the herbage at Jealott’s Hill was affected by an interaction between establishing seed mix and the number of years since plot establishment (seed × year: F6,171=11.6, p<0.001; Seed: F2,6=14.3, p<0.00; year: F3,171 =87.7, p<0.001; Fig. 67). Herbage phosphorus was in general slightly higher in the 2009 establishment year where legumes were included in the seed mix (‘grass & legume’ and ‘grass, legume & forb’) relative to where only grass was sown. However, there was a greater decline in herbage phosphorus for the former of these two treatments over the four year period from 2009-2012. Indeed, while herbage phosphorus remained relatively constant for the ‘grass’ only seed mixes at c. 0.19 % w/w, it collapsed from this same level to c. 0.11 % w/w and 0.15 % w/w respectively for the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes. It would seem that the inclusion of the non-legume forb component of the seed mix (‘grass, legume & forb’ plots) helped maintain phosphorus levels in the sward, but that the presence of legumes resulted in its depletion over a four year period at a rate much higher than would occur where only grasses were present. This overall decline in herbage phosphorus with time was also underpinned by a decline in soil phosphorus as already described above. Note that there was a slight peak in herbage phosphorus typical to all seed mixes in the drought year of 2010, which interestingly corresponded with a drop in Olsen’s Phosphorus in the same year. Although the mechanism underpinning this is not clear it would appear to be the result of a climatic rather than underlying treatment effect resulting in the greater deposition of phosphorus in drought stressed plant tissue. There was no significant effect of depth of seed bed cultivation (p>0.05). 0.3

0.25

0.2 2009 0.15 2010 0.1 2011 2012 Herbage Phosphorous (%) Phosphorous Herbage 0.05

0 Grass GL GLF Seed mix

Fig. 67. Effect of establishing seed mix and its interaction with year on the percentage of Phosphorus in the cut hay for Jealott’s Hill.

Herbage potassium (K): The percentage of potassium in the herbage cut from the swards at Jealott’s Hill responded to a significant interaction between establishing seed mixture, the depth of the cultivation practice used to establish their seed bed and the number of years since plot establishment (seed × cultivation × year: F15,126=9.13, p<0.001; seed: F2,6=34.1, p<0.001; cultivation: F2,33=1.76, p>0.05; year: F3,126=231.8, p<0.001; Fig. 68). For both seed mixes containing a sown legume component there was a decline in herbage potassium from c. 2.25 to 1.0 % w/w for the ‘grass & legume’ seed mix and c. 2.5 to 1.5 % w/w for the ‘grass, legume & forb’ seed mix. There seems to be some suggestion therefore that the inclusion of the non-legume forbs within the ‘grass,

88 legume & forb’ seed mix contributes to promoting the percentage of potassium within the herbage of the cut swards. While the initial levels in 2009 of herbage potassium were much higher for both of these seed mixes relative to the ‘grass’ only see mix (at around c. 1.75 % w/w) the rate of its decline in the ‘grass’ seed mix was much slower. By 2012 it had fallen to only c. 1.3 % w/w. In all cases, the collapse in herbage potassium between 2009 to 2012 for the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes is likely linked to the loss of legume dominance from the sward over this same time period. The relative maintenance of potassium in the sward of the ‘grass, legume & forb’ seed mix is likewise linked to the growing dominance of the non-legume forb component as the succession progressed for these plots. Although seed bed cultivation was part of this significant interaction between seed mix and year, its effects are not clear and are characterised by subtle changes in year to year rates in the overall decline in the percentage of herbage potassium.

a) ‘Grass’ seed mix 2.5

1.5 2009 2010 1 2011

0.5 2012 HerbagePotasium (% w/w)

0 Deep Shallow Cultivation b) ‘Grass & Legume’ seed mix. 3

2.5

2009 1.5 2010 1 2011 2012

HerbagePotasium (% w/w) 0.5

0 Deep Shallow Cultivation

89 c) ‘Grass, Legume & Forb’ seed mix. 3

2.5

2009 1.5 2010 1 2011 2012

HerbagePotasium (% w/w) 0.5

0 Deep Shallow Cultivation

Fig. 68. The effect of depth of seed bed cultivation over the four years of plot establishment on the3 percentage of potassium in cut herbage at Jealott’s Hill for the three seed mixtures. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Herbage calcium (Ca): The percentage of calcium in the cut herbage responded to an overall significant interaction between establishing seed mix, the timing of management (single or double yearly cuts), seed bed cultivation practice and the number of years since plot establishment (seed × timing × cultivation × year: F33,108=5.25, p<0.001; seed: F2,6=33.1, p<0.001; timing: F1,9=1.03, p>0.05; cultivation: F1,18=0.91, p>0.5; year: F3,108=51.1, p<0.001; Fig. 69). The ‘grass’ only seed mix typically produced a sward with lower percentage of herbage calcium than either the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes at c. 0.5 -0.6 % w/w. However, the calcium within the ‘grass’ only seed mix tended to maintain its percentage throughout the four year period from 2009 to 2012. For the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes the percentage calcium in the establishment year was typically much higher than that seen in the ‘grass’ seed mix, at around 0.8- 1.0 % w/w. however it tended to collapse over the four year period to end up at a similar or lower level to that found in the ‘grass ‘only plots. This rate of collapse was greatest for the ‘grass & legume’ seed mix, as the inclusion of the non-legume forb component of the sward seemed to buffer this decline in herbage calcium over the course of the succession. In addition, by 2012 the ‘grass & legume’ seed mix had swards with some of the lowest levels of calcium (c. 3 % w/w), while the ‘grass, legume & forb’ seed mixes remained comparable to that of the ‘grass’ only seed mix at c. 0.5 % calcium w/w. Again the effects of seed bed cultivation are not pronounced for any of the three seed mixes, and as such the interpretation of this treatment effect is not straightforward. There is some indication for the ‘grass, legume & forb’ seed mix that deep ploughing reduced the rate of decline in calcium from 2009-2012, a fact that may be linked to the better establishment of forbs under the deep ploughing treatment for the plots of this seed mix.

90 a) ‘Grass’ seed mix 1 0.9 0.8 0.7 0.6 0.5 0.4 2009 0.3 2010 0.2

HerbageCalcium w/w)(% 2012 0.1 2012 0 Deep Shallow Deep Shallow Rested Rested Typical Typical Timing of management and cultivation of seed bed b) ‘Grass & Legume’ seed mix 1 0.9 0.8 0.7 0.6 0.5 0.4 2009 0.3 2010 0.2

HerbageCalcium w/w)(% 2012 0.1 2012 0 Deep Shallow Deep Shallow Rested Rested Typical Typical Timing of management and cultivation of seed bed

c) ‘Grass, Legume & Forb’ seed mix 1.2

0.8

0.6 2009 0.4 2010

HerbageCalcium w/w)(% 0.2 2012 2012 0 Deep Shallow Deep Shallow Rested Rested Typical Typical Timing of management and cultivation of seed bed

Fig. 69. The effect on herbage calcium for the cut plots of Jealott’s Hill of establishing seed mix, the timing of management, the depth of seed bed cultivation and the years since plot establishment. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Herbage Magnesium (Mg): The percentage of magnesium in the cut herbage responded to an overall significant interaction between establishing seed mix, the timing of cutting management (single or double cuts yearly), seed bed cultivation depth and the number of years since plot establishment (seed × timing × cultivation × year: F33,108=5.02, p<0.001; seed: F2,6=21.8, p<0.001; timing: F1,9=0.45, p>0.05; cultivation: F1,18=5.57, p=0.02; year: F3,108=43.3, p<0.001; Fig. 70). Similar to the trend reported for herbage calcium, the percentage magnesium for the ‘grass’ only plots tended to remain relatively consistent over the four years between 2009 -2012 at c. 0.13 % w/w. However, the other two treatments showed clear declines in this mineral. Again the rate of decline was greatest for the ‘grass & legume’ seed mix falling from c. 0.2 to 0.1 % w/w magnesium by 2012, relative to a fall of 0.2 to 0.14 % w/w for the ‘grass, legume & forb’ seed mix over the same period. Also the rate of decline was higher for the ‘grass & legume’ seed mix, indicating that the presence of non-legume forbs as the succession progressed helped to buffer magnesium loss from herbage as the cover of legumes collapsed over time. There is again an indication similar to the effect seen for herbage calcium the ‘grass, legume & forb’ seed mix established by deep ploughing resulted in reduced rates of decline in this mineral from 2009-2012. This is lined to the better establishment of forbs under the deep ploughing treatment for the plots of this seed mix. a) ‘Grass’ seed mix 0.18 0.16 0.14 0.12 0.1

0.08 2009 0.06 2010 0.04 2012

HerbageMagnesium (% w/w) 0.02 2012 0 Deep Shallow Deep Shallow Rested Rested Typical Typical Timing of management and cultivation of seed bed b) ‘Grass & Legume’ seed mix 0.25

0.2

0.15

0.1 2009 2010

0.05 2012 HerbageMagnesium (% w/w) 2012 0 Deep Shallow Deep Shallow Rested Rested Typical Typical Timing of management and cultivation of seed bed

c) ‘Grass, Legume & Forb’ seed mix 0.3

0.25

0.2

0.15 2009 0.1 2010

0.05 2012 HerbageMagnesium (% w/w) 2012 0 Deep Shallow Deep Shallow Rested Rested Typical Typical Timing of management and cultivation of seed bed

Fig. 70. The effect on herbage magnesium in the cut plots of Jealott’s Hill of establishing seed mix, the timing of management, the depth of seed bed cultivation and the years since plot establishment. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Herbage sodium (Na): The percentage sodium within the cut herbage responded at Jealott’s Hill to a significant interaction between establishing seed mix and the number of years of plot establishment

(seed × year: F6,135=4.71, p<0.001; seed: F2,6=20.5, p<0.01; year: F3,135=28.8, p<0.001; Fig. 71). Initially at least the ‘grass’ only seed mix had swards with the lowest levels of herbage sodium, typically between 0.04 and 0.06 %. This tended to change little over the course of the four year succession in the plots. For the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes higher herbage sodium was found in the establishment year of 2009 at c. 0.09 % w/w, and while increasing slightly until 2011 it collapsed in 2012. This collapse in sodium for the cut herbage is linked with the large scale reduction in cover of legumes in both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes between 2011 and 2012. Again, it seems that the inclusion of the non-legume forbs in the ‘grass, legume & forb’ seed mix insulated this collapse in herbage sodium to some extent so that it did not fall below that of the ‘grass’ only plots at c. 0.05 % w/w. In contrast, the collapse in herbage sodium for the ‘grass & legume’ plots saw a fall to c. 0.02 % w/w by 2012, approximately half that of the other seed mixes in the same year.

0.14

0.12

0.1

0.08 2009 0.06 2010 2011 0.04 HerbageSodium (%) 2012 0.02

0 Grass GL GLF Seed mix

Fig. 71. The effect of the interaction between establishing seed mix and the number of years of plot establishment on the percentage sodium in the herbage of the cut plots at Jealott’s Hill.

The timing of the sward cuts (single or double cuts yearly) also had a slight effect on the percentage of sodium within the herbage, with plots under the rested management that received a single cut tending to have slightly less sodium by c. 0.02 % w/w (Timing: F1,11=5.58, p=0.04; Fig. 72). Although this effect is small, it is likely linked to shifts in the floral community composition of the repeatedly cut swards which were more likely to lost disturbance sensitive species that are characterised by higher leaf sodium concentrations. 0.09 0.08 0.07 0.06 0.05 0.04 0.03 HerbageSodium (%) 0.02 0.01 0 Typical Rested Timing of management

Fig. 72. The effect of the timing of cutting management on the percentage sodium in the herbage of the cut plots at Jealott’s Hill.

There was an overall significant effect of the depth of seed bed cultivation (F1,23=4.29, p=0.5; Fig. 73), with shallow cultivated plots having higher percentage sodium levels within the cut herbage. However, while significant this overall difference was very low, less than c. 0.02% w/w.

0.09 0.08 0.07 0.06 0.05 0.04 0.03 HerbageSodium (%) 0.02 0.01 0 Deep Shallow Cultivation

Fig. 73. The effect of the depth of seed bed cultivation on the percentage sodium in the herbage of the cut plots at Jealott’s Hill. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Herbage Dry matter: The dry matter of the cut herbage at Jealott’s Hill showed a significant response to the establishing seed mixture, the depth of cultivation method used to create its seed bed and the number of years of plot establishment (seed × cultivation × year: F15,126=5.57, p<0.001; seed: F2,6=28.5, p<0.001; Cultivation: F1,33=0.09, p>0.05; year: F3,126=226.3, p<0.001; Fig. 74). Typical to all seed mixes was that they tended to have herbage dry matter percentages of c. 92 %, and that this was generally consistent with the exception of a peak occurring in 2011. The origin for this peak is unclear, although it should be pointed out that it represents only a 1 – 1.5 % increase in the percentage of dry matter. It is possible this may be lined with the previous drought year in 2010, although there is no clear mechanism as to why this may be the case. Beyond this, differences resulting from cultivation depth and temporal trends of year were very minor and their interpretation from an applied perspective is not practical given the degree of between treatment variation. a) ‘Grass’ seed mix 94

93.5

92.5 2009 92 2010 91.5 2011 91 2012 Herbagedry matter (%w/w) 90.5

90 Deep Shallow Cultivation b) ‘Grass & Legume’ seed mix

93.5

92.5

92 2009 91.5 2010 2011 91 2012

Herbagedry matter (%w/w) 90.5

90 Deep Shallow Cultivation

c) ‘Grass, Legume & Forb’ seed mix 94

93.5

92.5 2009 92 2010 91.5 2011 91 2012 Herbagedry matter (%w/w) 90.5

90 Deep Shallow Cultivation

Fig. 74. The effect of the depth of establishing seed mix, the seed bed cultivation and the yeas since plot establishment on the percentage of dry matter in the herbage at Jealott’s Hill. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Herbage Pepsin cellulose digestibility: Herbage Pepsin cellulose digestibility of the cut sward at Jealott’s Hill was significantly influenced by an interaction between the establishing seed mixture, the timing of cutting management (single or double cuts) and the number of years since plot establishment (seed × timing × year: F15,126=4.52, p<0.001; seed: F2,6=7.80, p=0.02; timing: F1, 9=3.33, p>0.005; year: F3,126=616.5, p<0.001; Fig. 75). For all three seed mixes DODM remained relatively constant for the first three years of the succession (2009-2011) at 60-65 %. This then declined rapidly in the fourth year (2012) to 50% DODM for the ‘grass’ only seed mix, and around 45 % for the ‘grass & legume’ and ‘grass, legume & forb’ seed mix. Beyond this overarching trend there were only small differences between both the seed mixes and the timing of the cutting management. There is some indication that where legumes were a component of the seed mix under rested cutting (single sward cut) a more steady rate of decline in % DODM over the four years occurred when compared to the more rapid drop off between 2011 and 2012 for the typical cut plots (double sward cut). a) ‘Grass’ seed mix

40 2009

30 2010

(DODM%) 2011 20 2012 10

Herbage pepsin cellulose digetibility digetibility cellulosepepsinHerbage 0 Rested Typical Timing of management

b) ‘Grass & Legume’ seed mix 70

40 2009

30 2010

(DODM%) 2011 20 2012 10

Herbage pepsin cellulose digetibility digetibility cellulosepepsinHerbage 0 Rested Typical Timing of management

c) ‘Grass, Legume & Forb’ seed mix 80

50 2009 40 2010 30

(DODM%) 2011 20 2012 10

Herbage pepsin cellulose digetibility digetibility cellulosepepsinHerbage 0 Rested Typical Timing of management

Fig. 75. The effect of establishing seed mix, the timing of cutting management (rested vs. typical) and years since plot establishment on the herbage pepsin cellulose digestibility of the sward at Jealott’s Hill.

There was also a significant effect of the depth of seed bed cultivation in 2008 on DODM (F1,23=13.8, p=0.001; Fig. 76), and while this indicates that over the entire study period there was an increase in herbage digestibility under shallow cultivation, this effect difference was extremely small at c. 0.1 %. Again high variation and a small effect size suggest that the interpretation of such a difference has little biological meaning from an applied grassland management perspective.

61.5 61 60.5 60 59.5

59 (DODM (DODM %) 58.5 58

57.5 HerbagePepsin Cellulose Digestibility 57 Deep Shallow Cultivation

Fig. 76. The effect of seed bed preparation on the herbage pepsin cellulose digestibility (DODM) of the sward at Jealott’s Hill. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Animal performance (North Wyke Only)

Paddocks were grazed at the three herbage species levels for G (0.62 ha), GL (0.51 ha) or GLF (0.51 ha) plots on the four blocks and stocked with 12 groups of beef cattle from 11 June 2009 (following an initial cut across the whole of the experiment), 26 April 2010, 18 April 2011 and 16 May 2012. During the period that the Rested plots were excluded, the plot areas were temporarily reduced for G (0.51 ha), GL (0.41 ha) and GLF (0.41 ha). Resting occurred from 23 June to 8 September 2009, 18 June to 6 September 2010, 16 June to 12 September 2011 and 15 June to 10 September 2012 (Table 2b).

Livestock productivity at North Wyke was generally increased in 2010 compared with 2009, but this was largely due to 2009 being a shorter grazing season. Significantly fewer grazing days were supported on the unfertilised grass (G) than the GL and GLF plots in 2009 and 2012 but not in 2010 (Table 5). The increase in productivity from 2009 to 2010, as indicated by the grazing day total, of the grass only plots was probably largely influenced by the fact that the Existing grass sward, which was incorporated with the sown grass plots for grazing, was fertilised with 50 kg N per ha on 16 April 2010. Mean daily stocking rate was particularly high during June 2010 when the GL and GLF swards supported, on average, 15.5 cattle per ha, reflecting the period of peak growth of these swards.

Grazing in 2011 The North Wyke site was grazed in 2011 by Belgian Blue x Holstein Friesian steers born in August 2009 (mean 470 kg at turnout). Twelve groups of two steers were continuously stocked from turnout until 27 June and, for treatments G, GL and GLF, respectively daily live weight gains were

1.01, 1.04 and 1.06 kg per day (p = 0.867). Then the paddocks were rotationally grazed by two steers per block, spending 3.5 days per herbage species level paddock and had a mean daily live weight gain of 0.86 kg per day. The four groups were increased from two to four steers on 2 September and these grazed until they were removed on 17 October and in this period gained 1.18 kg per day on average.

Grazing in 2012 Twelve groups of three Hereford x Friesian Holstein cattle (2 steers and 1 heifer per group) born in September 2011 were continuously stocked (mean 224 kg at turnout) until 13 August when they were temporarily removed from the paddocks due to wet conditions. They returned on 10 September and grazed until 5 October when they were removed. For treatments G, GL and GLF respectively the mean daily live weight gains over the season were 0.78, 0.81 and 0.74 kg per day (p = 0.480).

The similarity in productivity between the grass legume and grass legume forb mixtures for all four years (Table 5) is a real effect, as these treatments were grazed separately. The inclusion of legumes (GL) or legumes and forbs (GLF) in the swards supported more grazing days and mean daily stocking rates in all four years (Table 5).

Cattle grazing days ha-1 Mean daily stocking rate cattle ha-1 2009 2010 2011 2012 2009 2010 2011 2012 G 425 992 394 502 3.8 5.9 2.2 3.5 GL 526 1235 481 612 4.7 7.2 2.6 4.3 GLF 526 1228 481 597 4.7 7.2 2.6 4.2 Fprobability: G v GL v GLF <0.001 0.093 - <0.001 <0.001 0.080 - <0.001 Table 5. North Wyke: Cattle grazing days and stocking rate.

Part 2. Further results not directly testing the seven key hypotheses.

The following section present results based on surveys undertaken at both sites, but do not provide conclusions that are directly related to the key seven hypotheses discussed in Part 1 above.

Soil chemistry

Statistical note for North Wyke: Analysis Part 1: Effects of seed mix and sward management on soil chemical properties. To test the effects of seed mix on soil chemical properties under either grazing or cutting management practises in a fully factorial manner. Analysis was done on the seed mix and management treatments and a sub set of the other treatments; only one level of timing of management (Typical) and cultivation (MC) was available so both of these were excluded from the analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management and Year.

Analysis Part 2: Effects of seed mix, cultivation and sward management on soil chemical properties. To test the effects of seed mix and seed bed preparation on soil chemical properties under either grazing or cutting management practises in a fully factorial manner. Analysis was done on the management and cultivation treatments and a sub set of the other treatments; two levels of seed mix (G & GLF) and one level of the timing of management (Typical) were available so timing of management was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Cultivation and Year

Total soil carbon

North Wyke: Analysis part 1) Total soil carbon declined between 2008 and 2012 (Year: F2,17=162.82, p<0.001)(Fig. 77). No significant effect of seed mix or management was found.

) 1

- 50

30 2008 20 2011

Total soil soil CarbonTotal kg (g 10 2012

0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 77. Effect of seed mix and sward management on soil total carbon at North Wyke

100

North Wyke: Analysis part 2) Total soil carbon was significantly higher in the minimally cultivated plots than the ploughed plots (Cult: F1,24=214.62, p =<0.001). There was a significant interaction between the effects of cultivation and year (Cult. x Year: F2,23=37.42, p=<0.001; year F2,23=33.53, p=<0.001) on total carbon at North Wyke. Total carbon concentrations declined throughout the duration of the experiment in the minimally cultivated plots, however in the ploughed plots concentrations tended to be more stable and generally increased from the establishment year except in the ‘grass only’ cut plots (Fig. 78). No significant seed mix or management treatment effects were found.

) 1 - 50

30 2008 20 2011 10 Total soil soil Carbon Total kg (g 2012

0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 78. Effect of seed mix, cultivation and sward management on soil total carbon at North Wyke

Jealott’s Hill (Total soil carbon): The total carbon from the 0-7.5 cm soil horizon at Jealott’s Hill was influenced by an interaction between the management used to maintain the sward and the number of years since plot establishment (management × year: F2,92=3.71, p=0.02; management: F1,11=0.07, p>0.05; year: F2,92=1.58, p>0.05; Fig. 79). The trend in total soil carbon was for it to decline from 2009 to 2012 where sward management was to cut for silage, while this trend was reversed under cattle grazing management. However, these trends showed a high degree of variation and the interpretation of these trends is hard from an applied management perspective. It is possible that the increase in soil carbon in the grazed plots is linked with accumulation of organic matter linked with cattle dung deposition, while the removal of vegetation after cutting similarly resulted in a decline in the organic matter accumulating in the soil. There was also a significant effect of the depth of cultivation practice used in the establishment of the seed bed in 2008, with shallow cultivation methods maintaining a slightly higher weight (c. 2 g kg-1) of carbon than deep ploughing

(cultivation: F1,23=23.8, p<0.001; Fig. 80). This positive effect of minimum cultivation techniques has been reported elsewhere and is in itself not an unexpected finding (Hutchinson et al., 2007; Liu et al., 2006). Tillage practices are known to have major effects on both the distribution of soil carbon as well as directly affecting the rates of organic matter decomposition (Liu et al., 2006). For this reason reduced tillage practices are most likely to maintain a relatively high soil carbon amount compared with conventional tillage with ploughing (Liu et al., 2006). No effect of seed mix or any higher order higher order interaction was found to influence soil carbon at Jealott’s Hill (p>0.05). Although soils represent a major store of carbon within the biosphere (Powlson et al., 2011), it has been suggested that the potential value of carbon sequestration within agricultural soils is relatively small in terms of its ability to mitigate against greenhouse gas emissions, representing perhaps 3-6% of fossil fuel emissions (Hutchinson et al., 2007). As there is unlikely to be a single simple solution to mitigate against greenhouse gas emissions, the contribution of minimum tillage approaches that promote

101 stores of carbon within the soil do represent an economically viable contribution which will confer benefits to society (Hutchinson et al., 2007).

29 1) - 27

25 2009 23 2010

21 2012 Total soil soil Carbin Total (g kg 19

17 Cut Grazed Management

Fig. 79. Effect of the interaction between sward management and the number of years since plot establishment on total soil Carbon at Jealott’s Hill.

28.5 )

1 28 - 27.5 27 26.5 26

Total soil soil Carbon Total (g kg 25.5 25 24.5 Deep Shallow Cultivation

Fig. 80. Effect of the depth of seed bed cultivation as undertaken in 2009 on the total soil carbon over the four year period. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Total soil nitrogen

North Wyke: Analysis part 1) There was a significant effect of year (Year: F2,17=74.5, p<0.001) on total soil nitrogen with concentrations generally decreasing throughout the duration of the experiment (Fig. 81). No significant effect of seed mix or management was found.

102

) 1 - 5

3 2008 2 2011

1 2012 Total soil soil NitrogenTotal kg (g 0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 81. Effect of seed mix and sward management on soil total nitrogen at North Wyke

North Wyke: Analysis part 2) There was a significant interaction between cultivation and year (Cult. x Year: F2,23=17.23, p=<0.001; Cult: F1,24=134.84, p=<0.001; Year: F2,23=19.69, p=<0.001) on total soil nitrogen at North Wyke. Total nitrogen concentrations were higher in the minimally cultivated plots than the ploughed plots (Fig. 82). In the minimally cultivated plots total nitrogen levels were highest in the establishment year whereas in the ploughed plots levels were similar in the first and final year of the experiment with the lowest concentrations found in 2011. No significant seed mix or management treatment effects were found.

) 1 - 5

3 2008 2 2011

1 2012 Total soil soil NitrogenTotal kg (g 0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 82. Effect of seed mix, cultivation and sward management on soil total nitrogen at North Wyke

Jealott’s Hill (Total soil nitrogen): At Jealott’s Hill total soil Nitrogen was found to be significantly higher where the depth of cultivation practice used in seed bed preparation in 2008 was shallow as opposed to deep ploughing (F1,43=6.65, p=0.01; Fig. 83). No other significant effects of either establishing seed mix, its subsequent management or a temporal component in terms of the number of years since margin establishment were found (p>0.05). This difference in total percentage nitrogen in the soil was small, with shallow cultivation supporting on average c. 1.5 g more Nitrogen per kg of soil. Tillage practices are known to have a major effect on the distribution of nitrogen (N) within the soil, as well as influencing N mineralization processes (Liu et al., 2006). Deep ploughing

103 has been shown to promote leaching of N from the soil in arable cropping systems (Aronsson & Stenberg, 2010; Rossella et al., 2007). Ploughing of grasslands increases soil organic N mineralization and has a consequent short term rise in soil mineral N (Velthof et al., 2010). Shallow cultivation to a depth of ca. 5 cm also represents a disturbance that can promote mineralization of organic N and cause an increase in soil mineral N amount, as reported by Velthof et al. (2010). The top horizons of soil have been shown to be the most responsive to cultivation practices in terms of effects on soil N content (Rossella et al., 2007). Nevertheless, minimum cultivation practices may be much less susceptible to leaching of N compared with deep ploughing. The drop in soil N in the top 7.5 cm of soil in the ploughed treatments at both sites compared with the minimally cultivated plots probably simply reflects the effect of burial of the organic matter richer surface soil and with it most of the soil N to below the sampling depth used in this study.

2.75

2.7

) 1 - 2.65

2.6

2.55

2.5

2.45 Total soil soil Total Nitrogen (g kg 2.4

2.35 Deep Shallow Cultivation

Fig. 83. Effect of the depth of initial seed bed cultivation on total soil Nitrogen over the four year sampling period at Jealott’s Hill. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Total soil phosphorus and Olsen phosphorus

North Wyke: Analysis part 1) To date no significant effects of either seed mix or management practise on soil total P or Olsen P have been observed. Total P content in the soil declined throughout the duration of the experiment (Year: F2,17=36.26, p<0.001) (Fig. 84). Soil Olsen P was highest in 2008 and lowest in 2011 (Year: F2,17=76.59, p<0.001) (Fig. 85).

104

2500

) 2000

1 -

1500

1000 2008 2011

Soil total (mg P Soil kg 500 2012

0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 84. Effect of seed mix and sward management on soil total phosphorus at North Wyke

25 )

1 20 -

10 2008 2011

Soil Olsen P Soil kg (mk 5 2012

0 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 85. Effect of seed mix and sward management on soil Olsen phosphorus at North Wyke

North Wyke: Analysis part 2) There was a significant interaction between cultivation and year on soil total phosphorus (Cult. x Year: F2,23=4.33, p=0.025; Cult: F1,24=5.94, p=0.023; Year: F2,23=36.73, p=<0.001) and on Olsen soil phosphorus (Cult. x Year: F2,23=12.51, p=<0.001; Cult: F1,24=10.58, p=0.004; Year: F2,23=50.27, p=<0.001) at North Wyke. Both total and Olsen P concentrations were higher in the minimally cultivated plots than the ploughed plots. Total P content in the soil declined throughout the duration of the experiment in the minimally cultivated plots and in the ploughed ‘grass only’ cut and ploughed ‘GLF’ grazed plots (Fig. 86). Soil Olsen P was highest in 2008 and lowest in 2011 in both the minimally cultivated and ploughed plots (Fig. 87). Total P levels were significantly affected by the management treatment (Man: F1,24=4.5.94, p=0.023) with higher concentrations found under the grazing regime compared with the cutting management. There was no significant management effect on Olsen P and no significant seed mix treatment effects were found for either Total P or Olsen P.

105

2000 1800

) 1600

1 - 1400 1200 1000 800 2008 600 2011

Soil total (mg P Soil kg 400 2012 200 0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 86. Effect of seed mix, cultivation and sward management on soil total phosphorus at North Wyke

25 )

1 20 -

10 2008 2011

Soil Olsen P Soil (mgkg 5 2012

0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 87. Effect of seed mix, cultivation and sward management on soil Olsen phosphorus at North Wyke

Jealott’s Hill (Total and Olsens Phosphorus): As was seen for total soil nitrogen, total phosphorus in the 0-7.5 cm soil horizon at Jealott’s Hill was found to respond significantly to the depth of cultivation for seed bed preparation in 2008 (F1,43=4.6, p=0.04; Fig. 88). Similar to the trend for nitrogen, total soil Phosphorus was on average highest where shallow cultivation methods were used, approximately 30 mg kg-1 higher than that seen under deep ploughing cultivation which had a mean soil Phosphorus of 655 mg kg-1. Overall soil phosphorus declined from 2009 to 2012 across all -1 plots, declining by c. 80-90 mg kg over this period (year: F2,94=65.8, p<0.001; Fig. 89). Shallow cultivation maintained higher soil phosphorus compared with ploughing, although this was not found in the case of Olsen Phosphorus (see below). This finding appears to be at odds with the findings of Omidi et al. (2008) who demonstrated that in the absence of tillage practices (in contrast to minimum tillage or conventional ploughing) a reduction of extractable soil phosphorus was seen. Neither the establishing seed mix nor the management of the sward was found to have a significant effect on total soil phosphorus.

106

710

)

1 )

- 700

1 -

g g g 690

gkg μ m 680 670 660 650 640

630 Total soil Phosphorous ( Phosphorous soil Total

Total soil Phophorus ( Phophorus soil Total 620 610 Deep Shallow Cultivation

Fig. 88. Change of total soil phosphorus across all treatments in response to the depth of cultivation practice used during seed bed preparation in 2008 at Jealott’s Hill. For seed bed cultivation, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

800

) 1

- 700

gkg

g g g m μ 600 500 400 300

200

)

Total soil Phophorus ( Phophorus soil Total 1 Total soil Phosphorous ( Phosphorous soil Total 100 0 2009 2010 2012 Year

Fig. 89. Change of total soil Phosphorus across all treatments during the succession of the grass swards at Jealott’s Hill.

Olsens Phosphorus at Jealott’s Hill differed from the response of total Phosphorus in that there was no identifiable effect of the depth of cultivation used in seed bed preparation. There was also no effect of either establishing seed mix or sward management (p>0.05). As for total soil Phosphorus,

Olsens Phosphorus responded to the age of the experimental plots (year: F2,138=5.35, p<0.01; Fig. 90) although the trend differed slightly. For Olsens Phosphorus there was evidence of a general overall decline from 2009 to 2012, however a slight increase in phosphorus (below the level of 2009) was identified in 2012.

107

18 )

1 16 -

g g kg 14 m 12 10 8 6 4

Olsens soil Olsens soil Phosphorus ( 2 0 2009 2010 2012 Year

Fig. 90. Change in Olsen’s soil phosphorus across all treatments during the succession of the swards at Jealott’s Hill.

Soil pH

North Wyke: Analysis part 1) Soil was slightly less acidic under the cutting management compared with the grazing management (Man: F2,17=5.38, p=0.032). Overall soil pH was highest in 2011 (Year: F2,17=6.9, p=0.006) and at its most acidic in the final year (2012) compared with the baseline soil samples collected in 2008 (Fig. 91). No significant effect of seed mix was found on soil pH. 6.2 6.1 6 5.9 5.8 5.7

5.6pH Soil 2008 5.5 5.4 2011 5.3 5.2 2012 MC MC MC MC MC MC G G GL GL GLF GLF cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical

Fig. 91. Effect of seed mix and sward management on soil pH at North Wyke

North Wyke: Analysis part 2) Soil was significantly less acidic in the ploughed plots than the minimally cultivated plots (Cult: F1,24=9.95, p=0.004) (Fig. 92). Year also had a significant effect (year. F2,23=14.17, p=<0.001) with soil pH tending to be highest in 2011. No significant seed mix or management treatment effects were found.

108

Soil pH Soil 3 2008 2011 2 2012 1

0 MC MC P P MC MC P P G G G G GLF GLF GLF GLF cut grazed cut grazed cut grazed cut grazed Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 92. Effect of seed mix, cultivation and sward management on soil pH at North Wyke

Jealott’s Hill (Soil pH): The response of soil pH in the 0-7.5 cm horizon at Jealott’s Hill showed some similarities to Olsens phosphorus in that the lowest value occurred half way though the succession of the exponential plots in 2010 (year: F2,138=12.54, p<0.001; Fig. 93). There was little difference in pH however at the start and end of the monitoring period with both soils in 2009 and 2012 having a similar pH of between 6.7-6.8, although there is some suggestion that it was slightly higher in 2012.

6.9

)

1 -

Soi 6.8 g kg g

l m 6.7 pH 6.6

6.5

6.4

6.3 Total soil Phosphorous ( Phosphorous soil Total 6.2 2009 2010 2012 Year

Fig. 93. Change in soil pH across all treatments during the succession of the grass swards at Jealott’s Hill.

109

Provision of summer and winter bird food resources

Statistical note for North Wyke:

Analysis Part 1: Effects of seed mix and sward management and timing of sward management on summer and winter bird food resources. To investigate the effects of seed mix, management and timing of management in a fully factorial manner on summer and winter bird food resources, analysis was done on the seed mix, management and timing of management treatments and a sub set of the cultivation treatment; only one level cultivation (MC) was available so cultivation was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Timing and Year.

Analysis Part 2: Effects of seed mix, cultivation and sward management on summer and winter bird food resources. To investigate the effects of seed mix, management and cultivation in a fully factorial manner on insect bird and insect pollinator food resources, analysis was done on the management and cultivation treatments and a sub set of the other treatments; two levels of seed mix (G) and (GLF) and one level of the timing of management (T) were available so timing was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Cultivation and Year.

Summer availability of sward active beetles

North Wyke: Analysis part 1) Following Saint-Germain et al (2007) the availability of bird food in the form of sward active beetles was analysed as total biomass as this provides a more accurate reflection of energy transfer through the food web to higher trophic levels than simple abundance. The mass of summer sward active beetles was affected by the interaction seed mix, subsequent sward management and its timing and the number of years since the plots were established (Seed x

Management x Timing x Year: F6,108=2.87, p=0.027; Seed: F2,6=29.75, p=<0.001; Man: F1,9=0.19, p=0.671; Timing: F1,18=19.29 p=<0.001; Year: F3,108=54.97 p=<0.001) (Fig. 94). There were also significant interactions between timing of management and year (Timing x Year: F3,108=8.55 p=<0.001) and management, timing of management and year (Man. x Timing x Year: F3,108=3.63 p=0.030) on beetle biomass. In all of the seed mixtures peak beetle biomass was found in 2010 and had generally collapsed by the final year (2012) and the ‘grass & legume’ sward supported the highest beetle biomass and the ‘grass only’ sward the least. Generally the ‘grass, legume & forb’ and ‘grass only’ swards supported higher beetle biomass in the rested plots than the typically managed plots. Differences between the timing of managements in the ‘grass & legume’ sward were negligible however generally higher beetle biomass was found in the typical cut plots than rested cut plots and the rested grazed plots supported higher beetle biomass than the typically managed grazed plots. In the establishment year (2009) all seed mixtures under both management regimes, beetle biomass was higher in the rested plots than the typically managed plots except in the grazed ‘grass & legume’ plots. In 2010 all typically cut swards supported higher beetle biomass than in the rested cut plots however more biomass was found in the rested grazed plots than the typically grazed plots. Although the management treatment was not significant there was an interaction between management and year (man. x year: F3,108=3.45 p=0.035) with generally more beetle biomass initially supported under the grazing management than the cutting management and from 2010 to 2012 the cutting regime supported more biomass.

110

140

120

100

2009 60 2010 2011

Total mass of beetles (mg)beetlesmass of Total 40 2012

Fig. 94. Effect of seed mix, sward management and timing of management on total beetle biomass at North Wyke.

North Wyke: Analysis part 2) Cultivation had an overall significant effect on beetle biomass (Cult:

F1,12=19.95, p=<0.001) with total beetle biomass higher in the ploughed plots than the minimally cultivated plots (Fig. 95). Seed mix also had an effect with higher beetle biomass in the ‘grass, legume & forb’ sward than the ‘grass only’ sward (Seed: F1,3=27.29, p=0.014). Although management was not significant there was an interaction between management and year (Man. X

Year: F3,72=4.50, p=0.013; Man: F1,6=0.09, p=0.769; year: F3,72=48.62, p=<0.001) with overall peak biomass found in 2010, higher biomass in the grazed plots in the establishment year and final year than the cut plots and the reverse trend in 2010 and 2011 with higher biomass in the cut plots. 20 18 16 14 12 10 2009 8 2010 6

4 2011 Beetle speciesBeetlerichness 2 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

111

Fig. 95. Effect of seed mix, sward management and cultivation on total beetle biomass at North Wyke.

Jealott’s Hill (Summer mass of sward active beetles): The mass of summer sward active beetles (as an index for overall invertebrate biomass within the sward) was directly affected by an interaction between the establishing seed mix, subsequent sward management, its timing and the number of years since the plots were established (seed × management × timing × year: F33,252=3.98, p<0.001; seed: F2,6=154.2, p<0.001; management: F1,9=46.1, p<0.001; timing: F1,18=81.1, p<0.001; year: F3,252=69.8, p<0.001; Fig. 96). The general trend across all three seed mixes was for rested management (single sward cut or suspended summer grazing) to support the highest biomass of beetles, particularly under cutting for the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes. For all seed mixes the peak in beetle biomass is clearly weighted towards the early establishment years, being highest in 2009 and largely having collapsed after four full years of establishment (2012). Although the most dramatic drop in biomass was between 2009 and 2010, under rested management biomass tended to decline less sharply over the four years. Finally there was an additional overall significant effect of the cultivation method used in initial seed bed preparation in

2008 (cultivation: F1,47=15.5, p<0.001; Fig. 97). Total mass of beetles active in the sward was on average 25 % higher where shallow cultivation methods had been sued to prepare the seed bed.

a) ‘Grass’ seed mix 50 45 40 35 30 25 2009 20 2010 15 2011

Total mass (mg) of beetles of (mg) mass Total 10 5 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

(b) ‘Grass & legume’ seed mix 250

200

150 2009 100 2010 2011

Total mass (mg) of beetles of (mg) mass Total 50 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing c) ‘Grass, legume & forb’ seed mixes

112

350

300

250

200 2009 150 2010

100 2011 Total mass (mg) of beetles of (mg) mass Total 50 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 96. The response of total mass of beetles at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

80 70 60 50 40 30 20

Total mass (mg) of beetles of (mg) mass Total 10 0 Deep Shallow Cultivation

Fig. 97. The response of total mass of beetles at Jealott’s Hill to cultivation method used to establish the seed mixes in 2008. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Winter grass seeds density

North Wyke: Analysis part 1) Winter seed data was log transformed to meet the assumptions of the ANOVA. The availability of winter grass seeds was affected by an interaction between sward management and timing of management (Man. x Timing: F1, 18=24.89, p=<0.001; Man: F1, 9=212.80, p=<0.001; Timing: F1, 18=14.43, p=0.001) and year and management (Year x Man: F3, 108=42.69, p=<0.001; Year: F3, 108=8.26, p=<0.001)( Fig. 98). Higher grass seed densities were found in the grazed plots compared with the cut plots in every year of the experiment, with the highest densities in 2009 and 2011. In the cut plots highest grass seed densities were in 2012 but this can be explained by the cut plots not receiving the second silage cut due to wet weather conditions. Grass seed density was highest in the grazed rested plots compared with the grazed typically managed plots whereas grass seed density was lower in the cut rested plots than the cut typically managed plots. Seed mix did not significantly affect the amount of available grass seed.

113

) 25

2 -

20 2009 15 2010

2011 Wintergrass seeds (m 10 2012

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut TypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRested

Fig. 98. Effect of seed mix, sward management and timing of management on winter grass seed density at North Wyke

North Wyke: Analysis part 2) Winter seed data was log transformed to meet the assumptions of the ANOVA. The availability of winter grass seeds was affected by an interaction between year and management (Year x Man: F3,72=16.03, p=<0.001; Year: F3,72=9.21, p=<0.001; Man: F1,6=41.52, p=<0.001 )(Fig. 99). Higher densities of grass seed were found in the grazed plots than the cut plots in all years. Seed bed preparation also had an effect on grass seed availability (Cult: F1,12=9.13, p=0.011) with generally higher grass seed abundance in the ploughed plots compared with the minimally cultivated plots. There was no clear pattern with year, however in the grazed plots highest grass seed densities were found in 2011 followed by 2009. Grass seed densities in the grass legume forb sown seed mix cut plots densities were highest in 2012 but this can be explained by the cut plots not receiving the second silage cut due to wet weather conditions. The seed mix treatment had no significant effect on grass seed density.

114

2) -

15 2009 10 2010

2011 Winter grass Winter seeds(m 5 2012

0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 99. Effect of seed mix, sward management and cultivation on winter grass seed density at North Wyke

Jealott’s Hill (Winter grass seed): The availability of winter grass seeds was affected by an interaction between the establishing seed mix, its sward management, the timing of this management and the number of years since the experimental plots were established (seed × management × timing × year: F33,300=9.09, p<0.001; seed: F2, 6= 3.91, p>0.05; management: F1, 9=7.82, p=0.02; timing: F1,18=39.3, p<0.001; year: F3,300=98.1, p<0.001; Fig. 100). The strongest temporal trend common to all three establishing seed mixes is a peak in grass seed density in the second establishment year (2010) under the rested timings of management (suspended summer grazing or single early cut). For the ‘grass’ seed mix the rested grazed plots have an exceptionally peak in grass seed density at c. 65 seed heads m-2, relative to a more common seed density of below 10 seed heads m-2 for other years and treatments. For the ‘grass and legume’ seed mix this pattern is reversed, with the rested cut plots in 2012 having the highest density of grass seed, although this remains considerably below the high seen in the ‘grass ‘ seed mix at c. 30 seed heads m-2. In the ‘grass, legume & forb’ seed mix seed head densities are even lower (c. 15 seed heads m-2) although the pattern remains with the rested grazed and cut plots supporting the highest densities in 2010 only, before collapsing in subsequent years.

115 a) ‘Grass’ seed mix

b) ‘Grass & Legume’ seed mix

c) ‘Grass. Legume & Forb’ seed mix

Fig. 100. The response of winter grass seed density at Jealott’s Hill in the (a) ‘grass), (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

116

Winter non-legume forb seed density

North Wyke: Analysis part 1) The density of non-legume forb seeds was dependent upon the interaction between sward management and seed mix (Man. x Seed: F2,9=11.38, p=0.003; Man: F1,9=20.98, p=<0.001; Seed: F2,6=11.06, p=0.010) and year and management (Man. x Year: F3,108=7.00, p=0.004; Year: F3,108=23.12, p=<0.001)(Fig. 101). Non-legume forb seed density declined throughout the duration of the experiment and density was highest in the grazed plots compared with the cut plots in all years. As expected non-legume forb seed density was highest in the grass legume forb treatments where forbs were sown. The timing of management treatment was not significant, however there was a significant interaction between timing of management and year (Year x Timing:

F3,108=7.72, p=0.002). In most instances typically managed plots supported higher forb seed densities than the rested plots, however the overall means showed the reverse (typical mean: 2.26m-2, rested mean: 3.11m-2) which can be accounted for by the minimally cultivated, grazed, ‘GLF’ seed mix, rested plot supporting the highest densities compared with any other treatments throughout the experiment.

) 50

2 -

30 2009

legumeforb seeds(m 2010 - 20 2011 2012

Winter nonWinter 10

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut TypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRested

Fig. 101. Effect of seed mix, sward management and timing of management on winter non-legume forb seed density at North Wyke

North Wyke: Analysis part 2) The density of non-legume forb seeds was dependent upon the interaction between year and management (Year x Man: F3,72=4.74, p=0.016; year: F3,72=22.56, p=<0.001; Man: F1,6=16.47, p=0.007 )(Fig. 102). Non-legume forb seed density declined throughout the duration of the experiment and more seeds were found in the grazed plots than the cut plots in all years. Cultivation had a significant effect on non-legume forb seed density (Cult: F1,12=13.94, p=0.003) with more seeds generally found in the ploughed treatments than the minimally cultivated treatments. There was a significant interaction between cultivation and seed mix (Cult. x Seed:

F1,12=18.99, p=<0.001) with more non-legume forb seeds in the ‘grass, legume & forb’ sward found in the ploughed plots than the minimally cultivated plots and in the ‘grass only’ sward more seeds were found where the seed bed had been prepared by minimal cultivation as opposed to ploughing. Seed

117 mix also affected seed density (F1,3=19.45, p=0.022), as expected more forb seeds were found in plots sown with the’ grass legume & forb’ seed mix than the ‘grass only’ seed mix.

)

2 - 50

30 2009

20 2010 legumeforb seeds(m - 2011 10 2012 0

Winter nonWinter P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 102. Effect of seed mix, sward management and cultivation on winter non-legume forb seed density at North Wyke

Jealott’s Hill (Winter non-legume forb seeds): The density of non-legume forbs at Jealott’s Hill was dependent on an interaction between the seed mix used to establish the experimental plots and the initial seed bed cultivation approach (seed × cultivation: F2,45=3.47, p=0.04; seed: F2,6=130.1, p<0.001; cultivation: F1,45=3.24, p>0.05; Fig. 103). As is to be expected, the highest density of non- legume forb seed heads was to be found in the ‘grass, legume & forb’ seed mix which represented the only plots that received direct sowing of forbs. In the ‘grass, legume & forb’ seed mix the density of non-legume forb seed heads was on average between 45 – 65 seed heads m-2, although this density was slightly higher where deep ploughing had been used as the initial seed bed cultivation practice in 2008. In addition, the density of non-legume forb seed heads was dependent on an interaction between the establishing seed mix, subsequent sward management, its timing and the number of years since the plots were established (seed × management × timing × year: F33,252=18.9, p<0.001; timing: F1,18=18.3, p<0.001; year: F3,252=53.5, p<0.001; Fig. 104). The temporal trend described for this relationship shows clear differences between those plots where forbs were part of the establishing seed mix and those where it was not. For the ‘grass, legume & forb’ seed mix there was a general trend of a peak in density in 2010, the second year of full plot establishment. This was most obvious where rested management occurred, particularly if the plot had been grazed. To a large part this is the result of the often persistent chicory stems which are not actively eaten by many cattle. In the rested plot the chicory also had an opportunity to develop and flower over the summer suspension of grazing. Under these circumstances seed head density could be extremely high (c. 400 seed heads of all forbs m-2), although as for grass seed head counts this collapsed to below 25 seed heads m-2 in all subsequent years. Where forbs were not part of the seed mix there was no peak in non-legume forb density in the second establishment year, rather a peak occurred after four years in 2012 for both the ‘grass’ and ‘grass & legume’ seed mixes. This reflects a gradual establishment of non- legume forbs into these experimental as a result of contamination from the surrounding more diverse swards. However, the density of non-legume forbs was for both ‘grass’ and ‘grass & legume’ seed mixes too low to be of any significance as bird food at typically less than 5 seed heads m-2.

118

90 80 70 60

)

2 -

40 Deep (m

30 Shallow legume forb seed head counts counts head seed forb legume - 20 10 0 Winter non Winter Grass GL GLF Seed mix

Fig. 103. Response of winter non-legume forb seeds at Jealott’s Hill to establishing seed mix and cultivation used in the preparation of the seed bed in 2008. a) ‘Grass’ seed mix

b) ‘Grass & Legume’ seed mix

1.8

)

2 - 1.6 1.4 1.2 1 2009 0.8 2010 0.6

0.4 2011

legume forb seed head (m seed head forb legume - 0.2 2012 0

Rest. Typi. Rest. Typi. Winter non Winter Cut Cut Graze Graze Management and its timing

119 c) ‘Grass, Legume & Forb’ seed mix

500

) 2 - 450 400 350 300 250 2009 200 2010 150 2011

legume forb seed head (m seed head forb legume 100 - 50 2012 0

Rest. Typi. Rest. Typi. Winter non Winter Cut Cut Graze Graze Management and its timing

Fig. 104. The response of winter non-legume forb seed head counts at Jealott’s Hill in the (a) ‘grass), (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

Winter legume forb seed density

North Wyke: Analysis part 1) Winter legume seed density was significantly affected by interactions between year and management (Year x Man: F3,108=32.27, p=<0.001; Year: F3,108=141.64, p=<0.001; Man: F1,9=44.90, p=<0.001), year and seed mix (Year x Seed: F6,108=36.54, p=<0.001; Seed: F2,6=140.99, p=<0.001), year, management and seed mix (Year x Man. x Seed mix: F6,108=8.17, p=<0.001) and year, management and timing of management (Year x Man. x Timing: F3,108=26.86, p=<0.001; Timing: F1,18=0.19, p=0.671)(Fig. 105). Legume seed density declined rapidly from the establishment year (2009), no legume seeds were recorded in the final two years. A higher density of legume seeds were found in the grazed plots compared with the cut plots in the ‘grass, legume & forb’ and ‘grass & legume’ swards in all years. In 2009 the ‘grass legume & forb’ seed mix supported a higher density of legume seeds than the ‘grass & legume’ seed mix, however the opposite was found in the second year (2010) though legume seed densities were extremely low in both of these swards by 2010. No legume seeds were recorded in the ‘grass only’ sward throughout the duration of the experiment. The timing of management treatment was not significant, however there was a significant interaction between timing of management and management (Man x Timing: F1,18=19.71, p=<0.001) with legume seed density in the grazed plots highest in the rested treatments compared with the typical treatments and in the cut plots the reverse was found although legume seed density was much lower in the cut plots.

120

)

2 - 25

20 2009 15 2010 10 2011

Winter legumeWinterseeds m 2012 5

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut TypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRestedTypicalRested

Fig. 105. Effect of seed mix, sward management and timing of management on winter legume seed density at North Wyke.

North Wyke: Analysis part 2) The density of legume forb seeds was dependent upon the interaction between year and seed mix (Year x Seed: F3,72=41.51, p=<0.001; Year: F3,72=61.28, p=<0.001; Seed: F1,3=61.69, p=0.004 )(Fig.106). Legume forb seed density declined throughout the duration of the experiment with no seeds recorded in the final two years of the experiment. As expected in both 2009 and 2010 more legume seeds were found in the plots sown with the ‘grass legume & forb’ seed mix than the ‘grass only’ seed mix. Cultivation had a significant effect on legume seed density (Cult:

F1,12=9.54, p=0.009) with higher seed abundance in the ploughed treatments than the minimally cultivated treatments. The management treatment had no significant effect on grass seed density.

) 2 - 10

6 2009

4 2010 2011 2 Winter legumeWinterseeds (m 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 106. Effect of seed mix, sward management and cultivation on winter legume seed density at North Wyke

Jealott’s Hill (Winter legume seeds): The density of legume seed heads at Jealott’s Hill was dependent on an interaction between the seed mix used to establish the experimental plots and the

121 initial seed bed cultivation approach (seed × cultivation: F2,45=4.86, p=0.01; seed: F2,6=17.5, p<0.01; cultivation: F1,45=1.56, p>0.05; Fig. 107). Again, the seed mixes which contained a legume component supported the highest density of winter legume seed heads, typically between 10-20 m- 2. Of these two seed mixes, the ‘grass & legume’ seed mix supported the highest legume seed density where deep cultivation was used to prepare the seed bed. In the case of the ‘grass, legume & forb’ seed mix there was no clear difference in the density of seeds in response to cultivation, although this was higher than that seen for the ‘grass’ only seed mix which had on average below two seed heads m-2. In addition to this response, the density of non-legume forb seed heads was dependent on an interaction between the establishing seed mix, subsequent sward management, its timing and the number of years since the plots were established (seed × management × timing × year: F33,252=3.98, p<0.001; management: F1,9=0.04, p>0.05; timing: F1,18=18.3, p<0.001; year: F3,252=11.2, p<0.001; Fig. 108). For the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes the peak in abundance was again associated with rested management timings (suspended summer grazing of a single early cut) with the peak in the second establishment year. In contrast to the patterns seen for grass and non- legume forb seed head densities described above, there was a greater propensity for this peak to extend beyond one year, most commonly into 2011. The highest peaks in legume seed head density were seen for the ‘grass & legume’ seed mix at c. 60 seed heads m-2. However, by 2012 legume seed head availability had collapsed to below 5 seed heads m-2 for all treatments combinations.

)

2 - 25

15 Deep 10 Shallow

5 Winter legume seed head counts (m counts head seedlegume Winter 0 Grass GL GLF Seed mix

Fig. 107. Response of winter legume seed head counts at Hill to the establishing seed mixture and the cultivation sued to prepare the seed bed in 2008. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

122

a) ‘Grass’ seed mix.

)

2 - 2.5

1.5 2009 2010 1 2011

0.5 2012 Winter legume seed head (m head seedlegume Winter 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing b) ‘Grass & Legume’ seed mix

120

)

2 - 100

60 2009 2010 40 2011

20 2012 Winter legume seed head (m head seedlegume Winter 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing c) ‘Grass, Legume & Forb’ seed mix

) 2 - 70 60 50 40 2009 30 2010 20 2011

10 2012 Winter legume seed head (m head seedlegume Winter 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 108. The response of winter legume seed head counts at Hill in the (a) ‘grass), (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

123

Plant community structure

Note see hypothesis 3 (H3) for non-legume forb and legume summed average percentage cover

Summed average percentage cover of grass

North Wyke: To investigate the effects of seed mix, management and timing of management in a fully factorial manner on the persistence of grass species, analysis was done on the seed mix, management and timing of management treatments and a sub set of the management; only one level of cultivation (MC) was available, therefore cultivation was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Timing and Year. The summed percentage cover of grass species was dependent upon the interaction between year and management (Year x Man: F3,108=4.63, p=0.008, Year: F3,108=30.91, p=<0.001, Man: F1,9=200.61, p=<0.001)(Fig. 109). Grass cover was highest in the first two years of the experiment (2009 mean: 62%, 2010 mean: 65.2%) and then cover declined (2011 mean: 46.1%, 2012 mean: 37.2%). Throughout the experiment grass cover was higher in the cut plots than the grazed plots. Timing of management did not significantly affect grass cover but there a significant interaction between management and timing of management (Man x Timing:

F1,18=15.08, p=0.001, timing of man: F1,18=0.21, p=0.815). Under the cutting regime grass cover was generally higher in the typical plots compared with the rested plots and the reverse trend was found under the grazing regime. . Seed mix had no significant effect on grass cover, however generally (the ‘grass & legume’ mix suppored the highest cover (mean: 56.5%) and surprisingly the ‘grass only’ mix the lowest cover (mean: 49.9%). 100 90 80 70 60 50 2009 40 2010 30 2011 20 2012 Summed grass percentage cover grasspercentage Summed 10 0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Rested Typical Rested Typical Rested Typical Rested Typical Rested Typical Rested

Fig. 109. Effect of seed mix, sward management and the timing of that management on the summed percentage cover of grass species at North Wyke

To investigate the effects of management and cultivation in a fully factorial manner on the persistence of grass species, analysis was done on the management and cultivation treatments and a sub set of the other treatments; only two levels of seed mix (G & GLF) and one level of the timing of

124 management (T) were available so timing of management was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Cultivation and Year. The summed percentage cover of grass species was dependent upon the interaction between year and cultivation (Year x Cultivation: F3,72=4.89, p=0.012; Year: F3,72=12.02, p=<0.001; Cult: F1,12=157.56, p=<0.001)(Fig. 110). Throughout the experiment grass cover was generally highest in the ploughed plots than the minimal cultivation plots regardless of sward type (Cult. x Seed: F1,12=27.63, p=<0.001). Grass cover was highest in the first two years of the experiment (2009 mean: 66.9%, 2010 mean: 70.5%) and then cover declined (2011 mean: 62%, 2012 mean: 49.8%). In all years grass cover was higher in the cut plots than the grazed plots (Year x Man:

F3,72=3.21, p=0.049; Man: F1,689.70, p=<0.001) and cut plots resulted in higher cover than the grazed plots regardless whether the seed bed was prepared by ploughing or minimal cultivation (Man x

Cult: F1,12=18.96, p=<0.001). Seed mix had no significant effect on grass cover, however the ‘grass only’ seed mix overall supported a higher cover than the ‘grass, legume & forb’ mix. 120

100

60 2009

40 2010 2011 20 2012

Summed grass Summedpercentage cover 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 110. Effect of seed mix, sward management and cultivation on the summed percentage cover of grass species at North Wyke

Jealott’s Hill (summed percentage cover of grasses): The summed percentage cover of grasses establishing into the swards at Jealott’s Hill was significantly influenced by an overall interaction between the establishing seed mix, its subsequent management, the cultivation method used in seed bed preparation and the number of years since the swards were established (Seed ×

Management × cultivation × year: F33,251=3.82, p<0.001; seed: F2,6=44.6, p<0.001; Management: F1,9=0.05, p>0.05; cultivation: F1,42=16.0, p<0.001; year: F3,251=52.3, p<0.001; Fig. 111). The overall temporal trend was for the summed percentage cover to increase over the four year period for all three seed mixes. However, the rate of this increase was far more extreme for both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes. In both of these seed mixes grass cover started at a relatively low at c. 20 % before increasing to 40-75% for the ‘grass & legume’ and 40-50 % for the ‘grass, legume & forb’ seed mix. This increase in the percentage cover of grasses in these two seed mixes over time is in part directly related to the collapse in the cover of legumes over the four years. Cutting management had a clear positive effect on grass cover over the four years for the ‘grass & legume’ seed mix, resulting in the highest increase in its percentage (an average of nearly 50%). A similar effect was also seen for the ‘grass, legume & forb’ seed mix although the difference between treatments was less clear. While the method used for seed bed cultivation was part of this overall significant interaction, its effects on the percentage cover of grasses are not pronounced. In addition to these effects, the timing with which management was applied resulted in an overall small but significant change in the summed percentage cover of grasses at Jealott’s Hill (timing:

125

F1,23=5.57, p=0.02; Fig. 112). The more intense typical management practice (either two sward cuts a year or continuous grazing without a summer rest period) benefited the grasses, so that there was a slight increase in their summed percentage cover, although by only c. 4 %. Overall the summed percentage cover of the grasses was typically quite high, at around % 50.

a) ‘Grass’ seed mix

b) ‘Grass & Legume’ seed mix

c) ‘Grass, Legume & Forb’ seed mix

126

Fig. 111. The response of the summed percentage cover of all grasses at Jealott’s Hill within the seed mixes of (a) ‘grass’ only, (b) ‘grass & legume’ and (c) ‘grass, legume and forb’ over the four years of the study (2009-2012) in response to management and initial seed bed cultivation technique. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

10 Grass sum percentage cover percentage sum Grass

0 Typical Rested Timing of management

Fig. 112. The effect of the timing of sward management at Jealott’s Hill on the summed percentage cover of grasses.

Plant community structure: species richness

North Wyke statistical note:

Analysis part 1: To investigate the effects of seed mix, management and timing of management in a fully factorial manner on sown grass, non-legume forb and legume forb species richness analysis was done on the management and timing of management treatments and a sub set of the other treatments; i. Grass species richness – all seed mix levels (G, GL &GLF) and one level of cultivation (MC). ii. Non-legume forb species richness – one seed mix level (GLF) and one level of cultivation (MC). iii. Legume species richness – two seed mix levels (GL & GLF) and one level of cultivation (MC).

Cultivation was therefore excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Management, Timing, Year and Seed mix for grass and legume species richness.

Analysis part 2: To investigate the effects of seed mix, management and cultivation in a fully factorial manner on the on sown grass, non-legume forb and legume forb species richness analysis was done on the management and cultivation treatments and a sub set of the other treatments; i. Grass species richness – all seed mix levels (G, GL &GLF) and one level of timing of management (typical).

127

ii. Non-legume forb species richness – one seed mix level (GLF) and one level of timing of management (typical). iii. Legume species richness – one seed mix level (GLF) and one level of timing of management (typical).

Timing of management was therefore excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Cultivation, Management, Year and Seed mix for grass and legume species richness.

Grass species richness

North Wyke: Analysis part 1) There was a significant interaction between management and year

(Man.x Year: F3,108=10.47, p=<0.001; Man. F1,9=2.12, p=0.179; Year F3,108=3.73, p=0.027) on grass species richness at North Wyke, although the management treatment was not significant (Fig. 113). Temporal changes are difficult to interpret with the overall grass species richness means gradually increasing year by year (2009: 2.958; 2010: 3.188; 2011: 3.292; 2012: 3.375). Grass species richness tended to increase under the cutting regime from the establishment to final year of the experiment whereas species richness tended to be more stable under the grazing management, the exception being in the ‘grass & legume’ seed mix plots where grass species diversity declined in 2010 and then increased in 2011. No significant seed mix or timing of management treatment effects were found. 5

4.5

3.5

2.5 2009

2 2010 2011 Grass speciesrichness Grass 1.5 2012 1

0.5

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Rested Typicl Rested Typical Rested Typical Rested Typical Rested Typical Rested

Fig. 113. Effect of seed mix, sward management and the timing of that management on grass species at richness North Wyke.

North Wyke: Analysis part 2) There was a significant interaction between cultivation and year (Cult. x Year: F3,72=6.09, p=0.004; Cult: F1,12=35.87, p=<0.001; Year: F3,72=7.76, p=<0.001) on grass species richness at North Wyke (Fig. 114). Temporal changes are difficult to interpret with the overall grass species richness means highest in the final year (2012), followed by the establishment year (2009)

128 and lowest in the middle years of the experiment (2009: 3.562; 2010: 3.312 2011: 3.312; 2012: 4.0). There was greater grass species diversity in the ploughed plots than the minimally cultivated plots in all years except in 2010 where the reverse trend was found, however in 2010 the number of grass species were generally similar in both seed bed preparation methods. No significant seed mix or management treatment effects were found. 6

3 2009

2 2010 2011 Grass speciesrichness Grass 1 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 114. Effect of seed mix, sward management and cultivation on grass species richness at North Wyke.

Jealott’s Hill (Grass species richness): Grass species richness at Jealott’s Hill was significantly affected by the interaction between establishing seed mix and the cultivation methods used in initial seed bed preparation (seed × cultivation: F2,80=6.85, p<0.01; Seed: F2,6=130.5, p<0.001; cultivation: F1,80=2.92, p>0.05; Fig. 115). Although the same species of grass were sown into all three seed mixtures, the ‘grass’ only seed mix supported the highest overall establishment of species. This high grass species richness in the ‘grass’ seed mix was not influenced by seed bed cultivation method, with no difference being apparent between the deep ploughed or shallow cultivation methods. This absence of a difference between the cultivation methods was also apparent for the ‘grass & legume’ seed mix, which while supporting on average 1 species less than the ‘grass’ only seed mix were affected by seed bed preparation. Only in the case of the ‘grass, legume & forb’ seed mix was there a clear differences between the cultivation methods, with the shallow cultivation supporting more species than the deep ploughing approach (although still fewer species that were established under the ‘grass’ only seed mix). It is likely that the greater establishment of the forb component of these seed mix where deep cultivation seed bed preparation was used (see below) resulted in some grass species being competitively excluded early on in sward establishment. The species richness of grasses that established in the swards were also affected by the interaction between sward management and year (management × year: F3,281=3.66, p=0.01; management: F1,11=0.18, p>0.05; year: F3,281=9.84, p<0.001; Fig. 116). However, the patterns are hard to interpret. At least for the cutting management there was a suggestion of a general trend of declining species richness from 2009 to 2011, although this was completely reversed by 2012 where grass species richness increased to a high of 3.5 species. It is possible that the initial good establishment of legumes from 2009 to 2011 was the cause of this initial trend of decreasing grass species richness, which was reversed by the general collapse in cover of at least some of the agricultural legume varieties in 2012 (e.g. T. pratense and T. hybridum). This collapse in legume cover in the final year of the study could have provided a less intense competitive environment allowing increased grass species colonisation. Where grazed management practices were applied to the sward this pattern is

129 less clear, although there is also a slight increase in grass species richness in 2012 which may similarly be attributed to a loss in the cover of some of the legume species. 4

3.5

2.5

2 Deep 1.5 Shallow

1 Grass sepeciesrichness Grass 0.5

0 Grass GL GLF Seed mix

Fig. 115. The effect on grass species richness of initial seed bed cultivation applied in the autumn of 2008 to the three seed mixes used to establish the swards. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Fig. 116. The response of grass species richness to grassland management and its interaction with years since establishment.

Non-legume forb species richness

North Wyke: Analysis part 1) There was a significant interaction between management and year

(Man. x Year: F3,36=6.05, p=0.013; Man: F1,3=0.79, p=0.439; Year: F3,36=32.80, p=<0.001) on non- legume species richness at North Wyke, although the management treatment was not significant (Fig. 117). Non-legume forb species richness declined throughout the duration of the experiment. From 2009 to 2011 non-legume forb diversity was higher in the grazed plots compared with the cut

130 plots, although this was reversed by 2012. Timing of management had no significant effect on non- legume forb species richness. 7

4 2009 3 2010

2 2011

legume forb species richnessforbspecies legume -

1 2012 Non 0 MC MC MC MC GLF GLF GLF GLF Grazed Grazed Cut Cut Typical Rested Typical Rested

Fig. 117. Effect of sward management and the timing of that management on non-legume forb species richness at North Wyke.

North Wyke: Analysis part 2) The species richness of non-legumes was significantly influenced by cultivation (Cult: F1,6=8.12, p=0.029) with more non-legume forb species found in the ploughed plots than the minimally cultivated plots (Fig. 118). Non-legume forb diversity declined throughout the duration of the experiment (Year: F3,36=68.01, p=<0.001). No significant management treatment effects were found, however there was a significant interaction between management and year

(Man. x Year: F3,36=9.90, p=0.001; Man: F1, 3=0.51, p=0.526) with non-legume forb species richness marginally higher in the grazed plots than the cut in the first three years with the reverse trend found in the final year (2012). 7 6 5 4 2009 3 2010 2 2011

1 legume forbspecieslegume richness - 2012

0 Non P MC P MC GLF GLF GLF GLF Grazed Grazed Cut Cut Typical Typical Typical Typical

Fig. 118. Effect of sward management and cultivation on non-legume forb species richness at North Wyke.

131

Jealott’s Hill (Non-legume forb species richness): The establishment of non-legume forbs at the Jealott’s Hill site was significantly affected by an interaction between the establishing seed mix, the subsequent management of the sward and the number of years since establishment occurred (seed

× management × year: F15,269=2.38, p<0.01; Seed: F2,6=400.1, p<0.001; Management: F1,9=5.16, p=0.05; year: F3,269=10.3, p<0.01; Fig. 119). The clearest overall effect, however, was of seed mix with the ‘grass, legume & forb’ seed mix unsurprisingly being characterised by supporting c. 4 additional species on average than either the ‘grass’ or ‘grass & legume’ seed mixes. It should be noted that for both of these latter two seed mixes there was a clear increase in non-legume forb species richness in 2012. In all cases this appears to be the result of gradual cross contamination between experimental plots as a result of animal and grass cutting activity spreading forb seeds into areas where they were not initially sown. The implications of management on non-legume forb species richness were less clear. There is some indication that for the ‘grass, legume & forb’ seed mix that by the final year of the study (2012) cutting management was more likely to support greater numbers of species (c. 1.5 more species) than plots that had been grazed. In addition to these treatment effects, there was an overall effect of seed bed cultivation. The species richness of non-legume forbs was significantly higher in plots where the seed bed had been established using the deep ploughing cultivation method as opposed to shallow cultivation

(Cultivation: F1,80=17.1, p<0.001; Fig. 120). However, cultivation showed no other significant interaction with any other of the treatment effects. The greater establishment of non-legume forb species into soil that had been deep ploughed in 2008 was in direct contrast to the response seen for grass species which showed greater establishment where the soil was shallow cultivated (see above).

3.5

2.5

1.5

1 legume forb species richness species forb legume

- 0.5 Non 0 Deep Shallow Cultivation

Fig. 119. The response of non-legume forb species richness to the cultivation method used in seed bed preparation in 2008. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

132

4 2009 3 2010

2 2011 legume forb species richness species forb legume - 1 2012

Non 0 Cut Graze Cut Graze Cut Graze Grass Grass GL GL GLF GLF Management and seed bed cultivation

Fig. 120. The response of non-legume forb species richness to establishing seed mix and its subsequent management over the four year duration of the study (2009 to 2012).

Legume forb species richness

North Wyke: Analysis part 1) The species richness of legumes was significantly influenced by the timing of management (Timing: F1,12=14.42, p=0.003) with more legume species found in the typically managed plots than the rested plots (Fig. 121). Legume diversity declined throughout the duration of the experiment (Year: F3,72=98.10, p=<0.001). No significant seed mix or management treatment effects were found. 4.5 4 3.5 3 2.5 2009 2 1.5 2010

1 2011 Legume species richnessspecies Legume 0.5 2012 0 MC MC MC MC MC MC MC MC GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Rested Typical Rested Typical Rested Typical Rested

Fig. 121. Effect of seed mix, sward management and the timing of that management on legume forb species richness at North Wyke.

North Wyke: Analysis part 2) The species richness of legumes was significantly influenced by cultivation (Cult: F1,6=44.70, p=<0.001) with more legume species found in the ploughed plots than the minimally cultivated plots (Fig. 122). Legume forb diversity declined throughout the duration of

133 the experiment (year: F3,36=75.06, p=<0.001) except in the ploughed cut plots where legume diversity was stable in the first two years. No significant management treatment effects were found on legume species diversity.

5 4.5 4 3.5 3 2.5 2009 2 2010 1.5

1 2011 Legumespeciesrichness 0.5 2012 0 P MC P MC GLF GLF GLF GLF Grazed Grazed Cut Cut Typical Typical Typical Typical

Fig. 122. Effect of sward management and cultivation on legume forb species richness at North Wyke.

Jealott’s Hill (legume species richness): The species richness of legumes establishing into the swards at Jealott’s Hill site were affected by an overall interaction between the establishing seed mix, its management and timing as well as the number of years since swards were established (seed

× management × timing × year: F33,299=10.6, p<0.001; Seed: F2,6=160.0, p<0.001; management: F1,9=3.97, p>0.05; year: F3,299=3.29, p=0.02; Fig. 123). The strongest overall temporal difference is between the seed mixes, with the ‘grass’ seed mix increasing in legume species richness while the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes tending to decline over time. These overall temporal trends occur independently of management and its timing. However, in the case of the grass only seed mix this increase in legume species richness over the four years is modest, being characterised by the establishment of T. repens and typically at most one other species such as T. pratense or T. hybridum. As already suggested these additional species are often the result of contamination though the activity of cattle or hay falling from cutting machinery after passing though some of the more diverse plots. The ‘grass’ seed mix rarely exceeds an average species richness of 1.5 legumes. For the ‘grass & legume’ seed mix the fall in legume species richness over the four years is greatest where grazing is used to manage the sward, particularly if typical grazing management is applied (i.e. no summer rest period). While typically 4-5 legume species are established in the ‘grass & legume’ seed mix, this fell to c. 2.5 species after four years in the typically grazed plots. This does however remain slightly higher under rested cutting (one cut per year) at c. 3.5 species. For the ‘grass, legume & forb’ seed mix the initial high species richness in 2009 is slightly lower than that occurring for the ‘grass & legume’ seed mix (c. 4.5 species). However, the same general pattern is seen in response to management and its timing as was seen for the more simple ‘grass & legume’ seed mix. In contrast to the species richness of both grasses and non-legume forbs, there was no effect of initial seed bed cultivation on the species richness of legumes (p<0.05).

134 a) ‘Grass’ seed mix

b) ‘Grass & Legume’ seed mix

c) ‘Grass Legume & forb ‘seed mix.

Fig. 123. The response of legume species richness for the grass (a), grass & legume (b) and grass, legume & forb (c) seed mixes to the interaction between management, its timing and the year of establishment (2009-2012).

135

Foraging resources for insect pollinators.

North Wyke statistical note: Analysis part 1: To investigate the effects of seed mix, management and timing of management in a fully factorial manner on insect bird and insect pollinator food resources, analysis was done on the seed mix, management and timing of management treatments and a sub set of the cultivation treatment; only one level cultivation (MC) was available so cultivation was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Timing and Year.

Analysis part 2: To investigate the effects of seed mix, management and cultivation in a fully factorial manner on insect bird and insect pollinator food resources, analysis was done on the management and cultivation treatments and a sub set of the other treatments; two levels of seed mix (G) and (GLF) and one level of the timing of management (T) were available so timing was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Cultivation and Year.

Non-legume forb flower counts

North Wyke: Analysis part 1) The density of non-legume forb flowers responded significantly to an interaction between seed mix and year (Seed x Year: F6,108=6.01, p=<0.001; Seed: F2,6=6.97, p=0.027; Year: F3,108=41.15, p=<0.001) and management and year (Man. X Year: F3,108=4.85, p=0.009; Man: F1,9=9.48, p=0.013)(Fig. 124). Peak non-legume forb flowers were found in 2010 followed by a decline in the final two years. In all years the ‘grass, legume & forb’ seed mix produced the highest number of non-legume forb flowers, comparable low counts were found in the ‘grass & legume’ and the ‘grass only’ seed mix. Higher densities of non-legume forb flowers were found in the grazed plots than the cut plots in all years except in the establishment year (2009) however flower numbers were so low the differences between the two treatments were negligible. Timing of management had no significant effect on non-legume forb densities.

136

30 2) - 25

2009 15 2010

2011 legume forb flower count (m countforbflower legume - 2012

10 Non

Fig. 124. Effect of seed mix, sward management and timing of management on non-legume forb flower counts at North Wyke.

North Wyke: Analysis part 2) Non-legume forb flower densities were affected by the interactions between cultivation, seed mix and year (Cult. x Seed x Year: F3,72=3.36, p=0.040; Cult: F1,12=28.65, p=<0.001; Seed: F1,3=13.38, p=0.035; Year: F3,172=51.86, p=<0.001), cultivation and year (Cult. x Year: F3,72=4.16, p=0.020) and seed mix and year (Seed x Year: F3,72=3.68, p=0.031)(Fig. 125). Peak non- legume forb flowers were found in 2010, one year after establishment and then flowers declined. In all four years the ploughed plots supported more forb flowers than the minimally cultivated plots in both the ‘grass, legume & forb’ and ‘grass only’ swards and unsurprisingly highest numbers of non- legume forb flowers were found in the ‘grass, legume & forb’ sward regardless of management

(Man. x Seed x Year: F3,72=3.23, p=0.046; Cult. x Seed x Man: F3,72=3.36, p=0.040). Although management was not significant there was an interaction between management and year (Man. x Year: F3,72=3.34, p=0.041; man: F1,6=0.00, p=0.987) with higher flower densities in the grazed plots than the cut plots in the establishment year and final year and the reverse trend in 2010 and 2011 with higher densities in the cut plots.

137

40 2) - 35 30 25 20 2009 15 2010 10 2011 5

2012 legume forbflowerlegume (m count - 0

Non P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 125. Effect of seed mix, sward management and cultivation on non-legume forb flower counts at North Wyke.

Jealott’s Hill (Non-legume forb flower counts pollinator feeding resource): The density of non- legume forb flowers, as a resource for insect pollinators, responded significantly to a higher order interaction between establishing seed mix, sward management, its timing of application and the number of years of establishment (seed × management × timing × year: F33,252=118.8, p<0.001; seed: F2,6=102.6, p<0.001; management: F1,9=82.4, p<0.001; timing: F1,18=25.2, p<0.001; year: F3,252=85.8, p<0.001; Fig. 126). Perhaps unsurprisingly, the density of non-legume forb flowers was highest where they had been a component of the initial seed mix, i.e. the ‘grass, legume & forb’ seed mix. In this seed mix, and in contrast to the pattern described below for legume flower, non-legume forbs tended to increase over the four year period so that their density was highest under by 2012. While this temporal trend for the ‘grass, legume & forb’ seed mix was consistent across all management types and their timing, the highest flower density were found where cutting as opposed to grazing was used. Where cutting management was used the density of non-legume forb flowers increased by a factor of four between 2009 to 2012. Different functional groups of pollinators tend to use different flower types, with bees often utilising legumes. In the initial establishment year there was little difference in the resources of legumes provided by the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes. However, it appears that the value of the ‘grass, legume & forb’ seed mix increased over the life span of the plots, so that the waning density of legume flowers over the four years (see below) was supplemented by the rise of non-legume forb flowers. Such non-legume forb flowers will be used by bees, but are of particular importance to hoverflies. In terms of creating an effective resource for pollinators the inclusion of non-legume forbs clearly has the potential to increase longevity. Non-legume forb flower counts were much lower in the ‘grass’ and ‘grass & legume’ seed mixes where they were never included in the establishing seed mix. However, in both cases there appears to have been something of a spike in density in the third year of full establishment (2011). This spike, seems to have been something of an anomaly and was not repeated in previous or subsequent year. There was also a significant interaction between establishing seed mix and the method used to provide a seed bed into which to sow those seed in 2009 (seed × cultivation: F2,45=9.78, p<0.001; Cultivation: F1,45=31.5, p<0.001; Fig. 127). As described above, the ‘grass, legume & forb’ seed mix supported the highest overall density of these flowers with evidence that their densities were highest where deep ploughing was used to establish the seed bed.

138 a) ‘Grass’ seed mix

) 2 - 0.9 0.8 0.7 0.6 0.5 2009 0.4 2010 0.3 2011 0.2

0.1 2012 legume forb flower counts (m counts flower legumeforb - 0

Non Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing b) ‘Grass & Legume’ seed mix

) 2 - 16 14 12 10 2009 8 2010 6 4 2011

2 2012 legume forb flower counts (m counts flower legumeforb - 0

Non Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

c) ‘Grass, Legume & Forb’ seed mix

)

2 - 20

15 2009 10 2010 2011 5

2012 legume forb flower counts (m counts flower legumeforb - 0

Non Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 126. The response of non-legume forb flower counts at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

139

) 2 - 8 7 6 5 4 Deep 3 Shallow

legume forb flower counts (m counts flower forb legume -

1 Non 0 Grass GL GLF Seed mix

Fig. 127. The response of non-legume forb flower counts as a resource for pollinators at Jealott’s Hill in response to establishing seed mix and the cultivation method used to establish them in 2008. For seed bed cultivation in 2008, deep = conventional ploughing to 25-20 cm, shallow = minimum tillage cultivation to c. 5cm depth.

Legume forb flower counts

North Wyke: Analysis part 1) Peak legume forb flowers were found in the establishment year and tended to decline in the following years with no legume flowers counted in the final year of the experiment (Fig. 128). Higher densities of legume flowers were found in the cut plots compared with the grazed plots in the first two years of the experiment (2009 & 2010) however slightly more legume flowers were found in the grazed plots in 2011 (Man. X Year: F3,108=10.16, p=<0.001; Man: F1,9=6.47, p=0.032; Year: F3,108=188.53, p=<0.001). In the establishment year highest numbers of legume flowers were present in the ‘grass, legume & forb’ swards and the least in the ‘grass only’ sward however more legume flowers were found in the ‘grass & legume’ sward than the ‘grass, legume & forb sward in 2010 and 2011 (Seed x Year: F6,108=44.83, p=<0.001; Seed: F2,6=670.89, p=<0.001). The rested plots supported higher numbers of legume flowers in 2009 than the typically managed plots however this trend was reversed in 2010 and 2011 with highest numbers found in the typically managed plots (Timing. x Year: F3,108=4.11, p=<0.020; Timing: F1,18=4.93, p=0.039). The density of legume forb flowers responded significantly to an interaction between management, seed mix and year (Man. x Seed x Year: F6,108=2.60, p=0.042), management, timing of management and year (Man. x Timing x Year: F3,108=3.38, p=0.039) and seed mix, timing of management and year (Seed x Timing x Year: F6,108=2.50, p=0.048). In 2009 the ‘grass, legume & forb’ seed mix supported the most legume flowers in both managements especially under the cutting regime, however in 2010 and 2011 the ‘grass & legume’ seed mix had the highest legume flower densities with more legume flowers found in the cutting management in 2010 and the grazing management in 2011. In 2009 more legume flowers were found in the cut typically managed plots then the cut rested plots and under the grazing management more legume flowers were found in the rested plots than the typically managed plots however in 2010 and 2011 more legume flowers were found in typically managed plots than the rested under both cut and grazing managements. In both the ‘grass, legume & forb’ and ‘grass & legume’ swards during the establishment year generally more legumes were

140 found in the rested plots than the typically managed plots with the reverse trend found in 2010 and 2011.

120

100

2) -

60 2009 2010 2011 40

2012 Legume flower counts (m countsflower Legume

0 MC MC MC MC MC MC MC MC MC MC MC MC G G G G GL GL GL GL GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Grazed Grazed Cut Cut -20 Typical Rested Typical Rested Typical Rested Typical Rested Typical Rested Typical Rested

Fig. 128. Effect of seed mix, sward management and timing of management on legume forb flower counts at North Wyke.

North Wyke: Analysis part 2) Seed bed preparation (Cult. x Year: F3,72=10.15, p=<0.001; Cult: F1,12=21.15, p=<0.001; Year: F3,72=56.78, p=<0.001) and seed mix (Seed x Year: F3,72=63.89, p=<0.001; Seed: F1,3=606.34, p=<0.001) had a significant effect on legume flower density (Fig. 129). Peak legume flowers were found in the establishment year and unsurprisingly a higher density of legume flowers were found in the ‘grass, legume & forb’ sward than the ‘grass only’ sward in all years. Regardless of sward type more legume flowers were found in the ploughed plots than the minimally cultivated plots (Cult. x Seed: F1,12=5.22, p=0.041). Although management was not significant there was an interaction between management and year (Man. x Year: F3,72=16.07, p=<0.001; Man: F1,6=2.49, p=0.0166) with higher legume flower densities in the cut plots in the first two years (2009 & 2010) and the opposing trend in the final two years (2011 & 2012) with higher densities in the grazed plots.

141

180

160 2) - 140 120 100 2009 80 60 2010 40 2011 20 2012 Legumeflower count(m 0 -20 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 129. Effect of seed mix, sward management and cultivation on legume forb flower counts at North Wyke.

Jealott’s Hill (Legume flower counts pollinator feeding resource): Legume flower counts, recorded as a foraging resource for insect pollinators, responded to a significant higher order interaction between establishing seed mix, sward management, its timing of application and year (seed × management × timing × year: F33,300=19.55, p<0.001; seed: F2,6=197.2, p<0.001; management: F1,9=62.6, p<0.001; timing: F1,18=1.02, p>0.05; year: F3,300=116.7, p<0.001; Fig. 130). For both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes there was a common pattern in the early years for the highest overall density of legume flowers to be found under cutting, as opposed to cattle grazing management. The density of flowers under cutting management was also highest where cutting management was rested and so was cut only once early in the year. For both the ‘grass & legume’ and ‘grass, legume & forb’ seed mixes there was clear evidence of a decline in the density of legume flowers over the four year sampling period. Significantly the density of legume flowers under cutting management, while starting off at a considerably higher level than that of grazed management, had by the fourth year fallen to densities lower than that seen in the grazed plots. This collapse in the density of legume flowers over the four year period was large, representing a fall from c. 70-90 flowers m-2 in 2009 to less than 5 flowers m-2 in 2012. In the ‘grass’ only seed mix, where legumes were not sown, the density of legumes was unsurprisingly consistently low in the initial establishment year, but as legumes became established over the four year period the density of legume flowers did increase. However, to never tended to exceed an average of 10 flower m-2. There was no significant effect of initial cultivation used in the establishment of the seed bed, either individually or as part of an interaction with the other treatments (p>0.05)

142

a) ‘Grass’ seed mix

) 2 - 8 7 6 5 2009 4 2010 3

2 2011 Legume flower counts (m counts flower Legume 1 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

b) ‘Grass & Legume’

100

) 2 - 90 80 70 60 50 2009 40 2010 30 2011

20 Legume flower counts (m counts flower Legume 10 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

c) ‘Grass, Legume & Forb’

) 2 - 80 70 60 50 2009 40 2010 30

20 2011 Legume flower counts (m counts flower Legume 10 2012 0 Rest. Typi. Rest. Typi. Cut Cut Graze Graze Management and its timing

Fig. 130. The response of legume flower counts at Jealott’s Hill in the (a) ‘grass’, (b) ‘grass & legume’ and (c) ‘grass, legume & forb’ seed mixes in response to sward management and the timing of that management from 2009 to 2012.

143

Invertebrate abundance and species richness

Additional analyses for North Wyke only

Statistical note for North Wyke: To investigate the effects of seed mix, management and cultivation in a fully factorial manner on abundance and diversity of pollen and nectar feeding invertebrates, analysis was done on the management and cultivation treatments and a sub set of the other treatments; two levels of seed mix (G) and (GLF) and one level of the timing of management (T) were available so timing was excluded from this analysis. This model therefore identifies the significant effects of all possible interactions between Seed Mix, Management, Cultivation and Year.

Note that as the experimental design use at Jealott’s Hill was balanced (i.e. all combinations of nested treatment levels were tested) single analyses was undertaken for all measures of invertebrate abundance and diversity. These have already been presented above in full under the experimental tests of Hypotheses 4 & 5.

Total pollinator abundance There was a significant interaction between seed mix, management and year (Seed x Man. X Year:

F3,72=5.41, p=0.006; Seed: F1,3=451.78, p=<0.001; Man: F1,6=17.28, p=0.006; Year: F3,72=17.48, p=<0.001 ) and between seed mix and year (Seed x Year: F3,72=23.79, p=<0.001) on total pollinator abundance (Fig.131). Generally highest numbers of pollinators were recorded in 2010 and the second highest in the establishment year (2009) with very low numbers found in 2011 and 2012. Overall the ‘grass, legume & forb’ sward supported more pollinators however as legume and non- legume forb percentage cover and flowers declined higher numbers of pollinators were recorded in the ‘grass only’ sward in the final two years of the experiment (2011 & 2012). The cut plots supported more pollinators than the grazed plots in both sward types. Although the cultivation treatment had no significant effect on total pollinator abundance there was an interaction between cultivation and year (Cult. x Year: F3,72=4.50, p=0.014) and cultivation and seed mix (Cult. x Seed: F1,12=4.95, p=0.046). Total pollinator abundance was overall higher in the ploughed than the minimally cultivated plots, however in the final two years of the experiment (2011 & 2012) slightly more pollinators were found in the minimally cultivated plots. Overall more pollinators were found in the ploughed ‘grass, legume & forb’ sward than the minimally cultivated ‘grass, legume & forb’ sward however the reverse trend was found in the ‘grass only’ sward with more pollinators in the minimally cultivated plots compared with the ploughed plots.

90 80 70 60 50 2009 40 30 2010 20 2011

Total pollinatorabundance Total 10 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

144

Fig. 131. Effect of seed mix, sward management and cultivation on total pollinator abundance at North Wyke.

Beetle abundance

Seed bed preparation had a significant effect on beetle abundance (Cult: F1,12=13.19, p=0.003) with generally higher numbers of beetles found in the ploughed plots than the minimally cultivated plots (Fig. 132). Peak beetle abundance occurred in 2010, followed by a decline in the following years and the ‘grass, legume & forb’ swards supported more beetles than the ‘grass only’ sward (Seed:

F1,3=33.57, p=0.010). There was an interaction between management and year (Man. x Year: F3,72=7.12, p=0.002; Man: F1,6=15.38, p=0.008; Year: F3,72=65.95, p=<0.001 ) with the grazed plots supporting more beetles than the cut plots in all years except in 2010.

90 80 70 60 50 2009 40 30 2010

Beetle abundanceBeetle 20 2011 10 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 132. Effect of seed mix, sward management and cultivation on beetle abundance at North Wyke.

Bumblebee abundance

There was a significant interaction between cultivation, seed mix and year (Cult. x Seed x Year:

F3,72=4.38, p=0.015; Cult: F1,12=4.83, p=0.048; Seed: F1,3=612.29, p=<0.001; Year: F3,72=44.59, p=<0.001), between seed mix, management and year (Seed x Man x Year: F3,72=5.48, p=0.006; Man: F1,6=24.58, p=0.003), between cultivation and seed mix (Cult x Seed: F1,12=10.78, p=0.007), between seed mix and management (Seed x Man: F1,6=14.45, p=0.009), between cultivation and year (Cult x Year: F3,72=6.85, p=0.002), between management and year (Man. x Year: F3,72=5.27, p=0.007) and between seed mix and year (Seed x Year: F3,72=53.51, p=<0.001) on bumblebee abundance (Fig. 133). Peak bumblebee abundance occurred in 2010 and then generally declined. Overall the ‘grass, legume & forb’ sward supported more bumblebees under both management types than the ‘grass only sward’ particularly in the ploughed plots. The cut plots supported more bumblebees than the grazed plots in all years, especially in the ‘grass, legume and forb’ sward.

145

70 60 50 40 2009 30 2010 20 2011 Bumblebeeabundance 10 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 133. Effect of seed mix, sward management and cultivation on bumblebee abundance at North Wyke.

Butterfly abundance

Management had a significant effect on butterfly abundance (Man: F1,6=19.99, p=0.004) with more butterflies found in the cut plots than the grazed plots (Fig. 134). Although seed mix and year had no individual significant effect on butterfly abundance there were some significant interactions;

(Seed x Year: F3,72=4.87, p=0.008; Seed: F1,3=2.47, p=0.214; Year: F3,72=1.65, p=0.197), (Seed x Man. x Year: F3,72=4.55, p=0.010) and (Seed x Man: F1,6=8.32, p=0.028). The ‘grass, legume & forb’ sward supported more butterflies than the ‘grass only’ sward in all years except the final year (2012). Overall the cut ‘grass, legume & forb’ sward supported more butterflies than the cut ‘grass only sward’ however the grazed ‘grass only’ sward supported slightly more butterflies than the grazed ‘grass, legume & forb’ sward. Cultivation had no significant effect on butterfly abundance. 9 8 7 6 5 2009 4 3 2010

Butterfly abundanceButterfly 2 2011 1 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 134. Effect of seed mix, sward management and cultivation on butterfly abundance at North Wyke.

146

Honeybee abundance

There was a significant interaction between cultivation and seed mix on honeybee abundance (Cult x

Seed: F1,12=8.63, p=0.012; Cult: F1,12=5.73, p=0.034; Seed: F1,3=34.54, p=0.010) with higher numbers of honeybees found in the ploughed plots than the minimally cultivated plots, particularly in the ‘grass, legume & forb’ sward (Fig. 135). There were also significant interactions between management and year (Man. x Year: F3,72=8.97, p=<0.001; Man: F1,6=15.02, p=0.008; Year: F3,72=12.52, p=<0.001), seed mix and year (Seed x Year: F3,72=14.79, p=<0.001) and seed mix, management and year (Seed x Man. X Year: F3,72=7.53, p=0.002). Generally honeybee abundance was highest in the establishment year (2009) and then declined with very few numbers found in the final two years of the experiment. Overall the cutting management supported more honeybees than the grazing regime especially in the ‘grass, legume and forb’ plots. 18 16 14 12 10 2009 8 6 2010

4 2011 Honeybee abundanceHoneybee 2 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 135. Effect of seed mix, sward management and cultivation on honeybee abundance at North Wyke.

Hoverfly abundance

Sward management had a significant effect on hoverfly abundance (Man: F1,6=10.27, p=0.019) with more hoverflies found in the cut plots than the grazed plots (Fig. 136). Overall lowest numbers of hoverflies were recorded in the establishment year (2009) with peak abundance in 2010 (Year:

F3,72=4.76, p=0.009). The seed mix and cultivation treatments had no significant effect on hoverfly abundance however there was a significant interaction between cultivation and year (Cult. x Year:

F3,72=3.42, p=0.034) and seed mix and year (Seed x Year: F3,72=4.76, p=0.009). In the first two years of the experiment (2009 & 2010) more hoverflies were found in the ‘grass, legume & forb’ sward than the ‘grass only’ sward and higher densities were found in the ploughed plots than the minimally cultivated plots however these trends were reversed in the final two years (2011 & 2012).

147

6 2009

4 2010

Hoverfly abundance Hoverfly 2011 2 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 136. Effect of seed mix, sward management and cultivation on hoverfly abundance at North Wyke.

Beetle species richness

Seed bed preparation had a significant effect on beetle diversity (Cult: F1,12=27.16, p=<0.001) with generally more species found in the ploughed plots than the minimally cultivated plots (Fig. 137).

Peak beetle species richness occurred in 2010 (Year: F3,72=52.72, p=<0.001), followed by a decline in the following years with similar low numbers of species found in the establishment year (2009) and the final year (2012). The ‘grass, legume & forb’ swards supported more beetle species than the

‘grass only’ sward (Seed: F1,3=29.77, p=0.012). Although the management treatment was not significant there was an interaction between management and year (Man. x Year: F3,72=7.88, p=0.003; Man: F1,6=1.28, p=0.301) with the grazed plots supporting more beetle species in 2009 and 2012 and the cut plots supporting more species in 2010 and 2011. 20 18 16 14 12 10 2009 8 2010 6

4 2011 Beetle speciesBeetlerichness 2 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 137. Effect of seed mix, sward management and cultivation on beetle species richness at North Wyke.

148

Bumblebee species richness

There was a significant interaction between seed mix and year (Seed x Year: F3,72=20.29, p=<0.001; Seed: F1,3=85.83, p=0.003; Year: F3,72=31.16, p=<0.001) with more bumblebee diversity found in the ‘grass, legume & forb’ sward than the ‘grass only’ sward in all years (Fig. 138). Bumblebee species richness was overall highest in the establishment year, closely followed by the second year (2010) before declining with no bumblebees found in the final year. Overall more bumblebee species were found in the cut plots than the grazed plots (Man: F1,6=11.07, p=0.016). Cultivation had no significant effect on bumblebee diversity. 4 3.5 3 2.5 2 2009 1.5 2010 1 2011 0.5 Bumblebeespeciesrichness 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 138. Effect of seed mix, sward management and cultivation on bumblebee species richness at North Wyke.

Butterfly species richness

Butterfly species richness was affected by the type of sward management (Man: F1,6=17.23, p=0.006) with more diversity found in the cut plots than the grazed plots (Fig. 139). Although seed mix was not significant there was a significant interaction between seed mix and management (Seed x Man.

F1,6=11.81, p=0.014; Seed: F1,3=2.01, p=0.252), seed mix, management and year (Seed x Man. x Year: F3,72=3.48, p=0.034; Year: F3,72=3.38, p=0.038) and seed mix and year (Seed x Year: F3,72=4.85, p=0.010). In the cut plots more butterfly species were recorded in the ‘grass, legume & forb’ sward than the ‘grass only’ sward in all years except 2012 whereas overall, more species were found in the grazed ‘grass only’ swards than the grazed ‘grass, legume & forb’ swards. Generally more butterfly species were recorded during the first two years of the experiment (2009 and 2010), particularly in the ‘grass, legume & forb’ swards. Species diversity declined in the ‘grass, legume & forb’ swards in 2011 with no butterflies recorded at all in 2012 whereas more species were recorded in the final two years than the first two years in the ‘grass only’ sward albeit diversity still remained very low in the ‘grass only’ swards. Cultivation did not significantly affect butterfly species diversity.

149

2.5

1.5 2009

1 2010 2011 0.5 Butterfly speciesButterflyrichness 2012 0 P MC P MC P MC P MC G G G G GLF GLF GLF GLF Grazed Grazed Cut Cut Grazed Grazed Cut Cut Typical Typical Typical Typical Typical Typical Typical Typical

Fig. 139. Effect of seed mix, sward management and cultivation on butterfly species richness at North Wyke.

Acknowledgements.

We wish to especially thank Richard Brand-Hardy at Defra and Steve Peel at Natural England. Also Ian Wilkinson of Cotswold Seeds Ltd for advice on the seed mixtures, and Innes McEwan as well as the farm staff at Syngenta Jealott’s Hill and North Wyke for assistance with managing the experiments. Thanks also to Lucy Hulmes, Sarah Hulmes, Jodey Peyton, Jo Savage, Markus Wagner and Bruce Griffith for help with field work, Pete Nuttall for the pollinator surveys and David Hogan for the North Wyke soil descriptions.

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Appendix 1a- l: Mean values (± SE) for all response variables from North Wyke including reference existing grassland plots managed as low input EK2 grasslands.

Appendix 1a: North Wyke soil chemistry

Total soil carbon (g kg-1)

Seed mix Man. Timing Cult. Means SE 2008 2011 2012 2008 2011 2012 Original grassland Cut Typical n/a 48.98 44.75 36.2 2.097 2.84 0.737 Graz. Typical n/a 51.08 47.27 39.61 1.188 3.53 1.899 Grass Cut Typical Min. Cult. 51.05 42.4 37.95 0.87 1.834 0.4 Cut Typical Plough 33.47 30.3 30.12 4.479 1.711 1.352 Graz. Typical Min. Cult. 49.73 48.83 37.88 1.798 2.671 2.577 Graz. Typical Plough 29.65 32.45 31.25 1.84 1.719 2.03 GL Cut Typical Min. Cult. 51.38 46.38 39.04 1.954 2.151 0.485 Graz. Typical Min. Cult. 52.27 48.1 40.56 1.438 2.702 2.469 GLF Cut Typical Min. Cult 49.93 45.77 37.46 0.861 1.696 0.44 Cut Typical Plough 29.62 31.3 30.95 1.394 1.418 2.273 Graz. Typical Min. Cult 50.22 45.35 39.86 0.709 0.729 1.174 Graz. Typical Plough 29.8 32.07 33.42 1.565 0.81 1.385

Total soil nitrogen (g kg-1)

Seed mix Man. Timing Cult. Means SE 2008 2011 2012 2008 2011 2012 Original grassland Cut Typical n/a 4.87 4.289 4.1 0.184 0.228 0.117 Graz. Typical n/a 4.857 4.538 4.321 0.211 0.304 0.214 Grass Cut Typical Min. Cult. 4.92 4.046 4.159 0.078 0.138 0.094 Cut Typical Plough 3.728 3.063 3.521 0.313 0.184 0.176 Graz. Typical Min. Cult. 4.96 4.694 4.265 0.147 0.249 0.255 Graz. Typical Plough 3.598 3.329 3.51 0.214 0.183 0.185 GL Cut Typical Min. Cult. 5.022 4.425 4.321 0.179 0.168 0.084 Graz. Typical Min. Cult. 5.062 4.569 4.434 0.199 0.271 0.289 GLF Cut Typical Min. Cult 4.957 4.345 4.258 0.086 0.179 0.068 Cut Typical Plough 3.475 3.272 3.606 0.189 0.163 0.238 Graz. Typical Min. Cult 4.92 4.348 4.358 0.026 0.109 0.130 Graz. Typical Plough 3.545 3.349 3.698 0.172 0.104 0.163

153

Soil Olsen’s extractable phosphorus (mg kg-1)

Seed mix Man. Timing Cult. Means SE 2008 2011 2012 2008 2011 2012 Original grassland Cut Typical n/a 17.25 5.826 9.171 1.163 2.328 0.932 Graz. Typical n/a 14.69 7.649 9.69 1.021 1.433 0.749 Grass Cut Typical Min. Cult. 17.9 6.789 10.8 0.286 2.541 0.779 Cut Typical Plough 10.56 3.184 7.83 2.594 1.253 1.183 Graz. Typical Min. Cult. 17.5 9.879 10.67 1.797 2.25 2.835 Graz. Typical Plough 8.54 3.769 8.46 0.867 1.721 0.972 GL Cut Typical Min. Cult. 17.25 7.988 8.79 0.495 1.477 0.632 Graz. Typical Min. Cult. 16.89 8.542 11.33 0.503 3.208 0.683 GLF Cut Typical Min. Cult 17.84 9.621 9.91 2.269 2.294 1.279 Cut Typical Plough 8.48 5.378 6.1 1.447 2.574 1.295 Graz. Typical Min. Cult 18.71 7.316 10.55 3.822 3.118 1.459 Graz. Typical Plough 9.48 2.664 9.34 1.424 1.655 1.351

Total soil phosphorus (mg kg-1)

Seed mix Man. Timing Cult. Means SE 2008 2011 2012 2008 2011 2012 Original grassland Cut Typical n/a 1072 1068 858.5 54.02 47.23 35.63 Graz. Typical n/a 1035 1092 868.3 55.56 95.87 32.97 Grass Cut Typical Min. Cult. 1108 1006 838.7 113.01 55.31 41.3 Cut Typical Plough 900 820 796.6 38.34 26.37 41.44 Graz. Typical Min. Cult. 1234 1085 941.1 70.78 57.6 72.11 Graz. Typical Plough 879 967 782.1 50.75 40.04 27.35 GL Cut Typical Min. Cult. 1103 1051 885.2 40.88 42.17 36.91 Graz. Typical Min. Cult. 1088 1138 943.7 44.1 123.63 41.34 GLF Cut Typical Min. Cult 1178 1027 905.2 67.36 106.19 44.28 Cut Typical Plough 807 852 764.5 55.31 40.69 32.15 Graz. Typical Min. Cult 1532 1057 899 338.49 30.64 24.0 Graz. Typical Plough 999 963 874.6 40.21 39.88 46.11

Soil pH

Seed mix Man. Timing Cult. Means SE 2008 2011 2012 2008 2011 2012 Original grassland Cut Typical n/a 5.75 5.89 5.81 0.05 0.11 0.01 Graz. Typical n/a 5.68 5.79 5.63 0.08 0.11 0.02 Grass Cut Typical Min. Cult. 5.82 5.86 5.79 0.06 0.11 0.06 Cut Typical Plough 5.92 6.36 6.02 0.08 0.24 0.11 Graz. Typical Min. Cult. 5.72 5.76 5.60 0.11 0.12 0.02 Graz. Typical Plough 5.85 5.66 5.78 0.07 0.16 0.08 GL Cut Typical Min. Cult. 5.71 5.97 5.73 0.08 0.10 0.06 Graz. Typical Min. Cult. 5.76 5.79 5.62 0.11 0.14 0.06 GLF Cut Typical Min. Cult 5.66 5.87 5.69 0.10 0.11 0.10 Cut Typical Plough 5.99 6.07 5.82 0.08 0.08 0.09 Graz. Typical Min. Cult 5.81 5.69 5.66 0.08 0.15 0.03 Graz. Typical Plough 5.91 5.99 5.73 0.04 0.06 0.07

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Appendix 1b: North Wyke soil structure

Soil bulk density 0-10cm (g cm3)

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original grassland Cut Typical n/a 0.76 0.79 0.73 0.05 0.03 0.01 Graz. Typical n/a 0.70 0.78 0.76 0.01 0.03 0.01 Grass Cut Typical Min. Cult. 0.77 0.76 0.68 0.06 0.04 0.02 Cut Typical Plough 0.77 0.86 0.86 0.01 0.05 0.01 Graz. Typical Min. Cult. 0.72 0.76 0.74 0.02 0.04 0.01 Graz. Typical Plough 0.78 0.85 0.84 0.03 0.04 0.01 GL Cut Typical Min. Cult. 0.72 0.78 0.74 0.02 0.05 0.02 Graz. Typical Min. Cult. 0.68 0.75 0.75 0.02 0.40 0.01 GLF Cut Typical Min. Cult 0.79 0.81 0.77 0.02 1.04 0.01 Cut Typical Plough 0.82 0.86 0.90 0.01 0.04 0.01 Graz. Typical Min. Cult 0.69 0.78 0.77 0.01 0.04 0.01 Graz. Typical Plough 0.77 0.85 0.82 0.04 0.03 0.02

Soil bulk density 10-20cm (g cm3)

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original grassland Cut Typical n/a 1.01 1.04 1.07 0.11 0.04 0.01 Graz. Typical n/a 0.92 1.01 1.02 0.02 0.04 0.02 Grass Cut Typical Min. Cult. 1.02 1.03 1.03 0.09 0.05 0.02 Cut Typical Plough 0.86 0.94 0.94 0.04 0.04 0.01 Graz. Typical Min. Cult. 0.90 0.99 1.01 0.02 0.03 0.01 Graz. Typical Plough 0.82 0.92 0.87 0.03 0.04 0.03 GL Cut Typical Min. Cult. 0.90 1.00 1.06 0.04 0.05 0.02 Graz. Typical Min. Cult. 0.88 0.98 1.00 0.03 0.06 0.01 GLF Cut Typical Min. Cult 0.99 1.04 1.07 0.03 0.05 0.01 Cut Typical Plough 0.87 0.91 0.94 0.01 0.04 0.02 Graz. Typical Min. Cult 0.90 0.98 1.02 0.0 0.04 0.02 Graz. Typical Plough 0.81 0.87 0.91 0.2 0.04 0.01

155

Appendix 1c: North Wyke soil nutrient loss

Total oxidised nitrogen (mg l-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2012 2012 Original Cut Typical n/a 0.06 0.12 0.15 0.05 0.03 0.02 0.06 0.02 grassland Graz. Typical n/a 0.02 0.09 0.08 0.04 0.00 0.00 0.00 0.01 Grass Cut Typical Min. Cult 0.02 0.09 0.07 0.05 0.00 0.00 0.00 0.03 Cut Typical Plough 0.08 0.09 0.12 0.03 0.06 0.00 0.07 0.01 Graz. Typical Min. Cult 0.02 0.09 0.08 0.08 0.00 0.00 0.01 0.03 Graz. Typical Plough 0.03 0.10 0.07 0.03 0.01 0.01 0.00 0.00 GL Cut Typical Min. Cult 0.05 0.09 0.08 0.02 0.03 0.00 0.00 0.00 Graz. Typical Min. Cult 0.02 0.17 0.08 0.02 0.01 0.05 0.01 0.00 GLF Cut Typical Min. Cult 0.22 0.09 0.13 0.06 0.19 0.00 0.04 0.03 Cut Typical Plough 0.33 0.15 0.28 0.10 0.26 0.06 0.19 0.04 Graz. Typical Min. Cult 0.02 0.09 0.11 0.03 0.00 0.00 0.03 0.02 Graz. Typical Plough 0.03 0.09 0.09 0.02 0.00 0.00 0.02 0.00

Total phosphorus (µg l-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 34.03 30.14 22.85 0 21.2 5.49 5.82 grassland Graz. Typical n/a 4.13 21.63 13.1 154.2 3.33 6.33 2.28 82.3 Grass Cut Typical Min. C 1.05 25.92 15.03 29.04 0.61 9.81 1.34 6.92 Cut Typical Plough 0.55 14.16 11.77 20.68 0.48 6.06 2.24 2.86 Graz. Typical Min. C 2.74 20.84 16.63 41.93 0.95 3.00 4.91 13.8 Graz. Typical Plough 1.30 15.24 16.6 21.86 0.83 0.92 2.31 0.52 GL Cut Typical Min. C 2.83 12.35 16.54 35.26 1.54 0.86 2.98 11.1 Graz. Typical Min. C 8.18 62.03 16.74 21.41 2.35 13.01 3.23 3.55 GLF Cut Typical Min. C 4.32 20.17 95.84 31.82 2.39 3.95 81.82 9.89 Cut Typical Plough 1.26 41.52 13.47 36.85 0.82 13.47 2.27 16.7 Graz. Typical Min. 1.79 20.77 52.08 76.37 0.71 2.81 18.85 20.2 Graz. Typical Plough 1.12 20.58 22.1 35.00 0.65 5.00 2.54 5.74

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Appendix 1d: North Wyke herbage dry matter production from combined yearly silage cuts (tonnes ha-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 7.39 7.81 9.18 10.47 0.19 0.53 0.68 1.55 grassland Grass Cut Typical Min. C 6.02 6.25 8.08 8.36 0.70 0.06 0.47 0.71 Cut Typical Plough 7.02 5.35 6.18 6.62 0.67 0.42 0.51 0.53 Cut Rested Min. C 5.16 6.25 7.22 8.45 0.57 0.18 0.86 0.54 GL Cut Typical Min. C 7.44 7.74 8.07 8.00 0.65 0.60 0.60 0.20 Cut Rested Min. C 7.57 7.34 8.27 9.5 0.48 0.73 0.65 0.27 GLF Cut Typical Min. C 7.87 7.03 7.07 8.21 0.48 1.32 1.05 0.70 Cut Typical Plough 11.70 8.1 7.29 7.28 0.83 0.97 1.33 0.20 Cut Rested Min. C 6.92 5.86 6.38 8.67 0.46 1.13 1.0 0.70

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Appendix 1e: North Wyke herbage forage quality

Herbage nitrogen (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 16.93 17.6 16.49 15.91 1.36 0.95 0.81 1.44 Graz. Typical n/a 28.46 20.45 20.07 25.27 1.50 1.12 0.52 1.8 Grass Cut Typical Min. C 12.94 14.4 13.76 14.03 0.48 0.39 0.23 1.23 Cut Typical Plough 11.61 13.51 13.7 11.98 0.26 0.33 0.51 1.77 Cut Rested Min. C 15.09 13.71 13.24 16.47 0.81 0.29 0.45 2.21 Graz. Typical Min. C 27.25 18.89 18.8 24.24 1.27 1.04 1.13 1.70 Graz. Typical Plough 22.18 17.67 19.06 22.19 0.91 0.97 1.31 1.71 Graz. Rested Min. C 23.55 18.41 18.48 30.45 0.35 1.15 0.82 3.06 GL Cut Typical Min. C 17.27 15.84 15.5 14.31 1.47 0.83 1.17 1.4 Cut Rested Min. C 17.61 15.86 14.05 16.29 1.53 0.43 0.24 2.70 Graz. Typical Min. C 34.95 20.64 20.59 24.15 1.78 1.39 1.52 2.35 Graz. Rested Min C 37.04 18.92 18.83 28.25 0.29 0.92 0.55 3.20 GLF Cut Typical Min. C 16.74 16.02 14.71 16.58 1.28 0.40 0.72 2.27 Cut Typical Plough 16.46 17.24 15.39 14.91 1.61 0.73 0.99 1.86 Cut Rested Min. C 16.49 16.16 15.9 16.79 1.57 0.40 0.97 2.79 Graz. Typical Min. C 33.14 20.05 20.81 24.91 1.58 0.71 1.70 1.63 Graz. Typical Plough 29.75 24.8 23.14 25.2 1.59 1.52 2.02 1.75 Graz. Rested Min.C 36.02 19.38 18.46 28.11 1.25 0.87 0.76 2.86

Herbage phosphorus (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 2.65 2.89 2.54 1.88 0.15 0.12 0.13 0.14 Graz. Typical n/a 3.7 2.8 2.8 2.4 0.04 0.10 0.10 0.18 Grass Cut Typical Min. C 2.31 2.58 2.46 1.74 0.10 0.10 0.10 0.12 Cut Typical Plough 1.86 2.34 2.49 1.43 0.09 0.07 0.15 0.20 Cut Rested Min. C 2.35 2.41 2.28 1.85 0.11 0.07 0.07 0.18 Graz. Typical Min. C 3.78 2.75 2.84 2.46 0.13 0.11 0.12 0.15 Graz. Typical Plough 2.88 2.64 2.73 2.15 0.34 0.07 0.14 0.13 Graz. Rested Min. C 3.35 2.65 2.69 2.76 0.16 0.07 0.09 0.21 GL Cut Typical Min. C 2.41 2.33 2.49 1.78 0.05 0.05 0.17 0.16 Cut Rested Min. C 2.31 2.51 2.2 1.88 0.15 0.04 0.07 0.19 Graz. Typical Min. C 3.23 2.84 2.96 2.63 0.12 0.09 0.10 0.18 Graz. Rested Min. C 3.35 2.7 2.81 2.69 0.20 0.11 0.08 0.20 GLF Cut Typical Min. C 2.29 2.45 2.48 2.09 0.10 0.12 0.15 0.21 Cut Typical Plough 2.13 2.29 2.3 1.85 0.07 0.06 0.13 0.22 Cut Rested Min. C 2.26 2.5 2.43 1.88 0.11 0.13 0.16 0.24 Graz. Typical Min. C 3.38 2.74 2.83 2.56 0.22 0.10 0.16 0.15 Graz. Typical Plough 3.23 2.86 2.86 2.28 0.03 0.17 0.11 0.09 Graz. Rested Min. C 3.18 2.8 2.7 2.56 0.14 0.13 0.12 0.16

158

Herbage calcium (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 7.03 6.06 7.03 4.95 0.72 0.34 0.31 0.40 Graz. Typical n/a 6.46 6.02 6.61 5.2 0.37 0.23 0.42 0.15 Grass Cut Typical Min. C 5.16 5.50 6.21 4.8 0.36 0.35 0.20 0.34 Cut Typical Plough 4.60 5.47 6.03 3.76 0.29 0.70 0.38 0.31 Cut Rested Min. C 4.93 5.17 6.19 4.81 0.20 0.25 0.21 0.34 Graz. Typical Min. C 6.38 5.42 5.96 5.1 0.31 0.26 0.24 0.15 Graz. Typical Plough 5.29 5.13 5.55 5.01 0.31 0.44 0.26 0.20 Graz. Rested Min. C 6.34 4.87 5.44 4.46 0.28 0.24 0.14 0.20 GL Cut Typical Min. C 7.93 7.00 7.75 4.7 1.42 0.94 1.00 0.31 Cut Rested Min. C 6.95 5.88 6.11 4.49 1.05 0.46 0.23 0.41 Graz. Typical Min. C 9.27 6.18 6.99 5.2 0.59 0.49 0.76 0.16 Graz. Rested Min. C 13.77 5.18 5.49 4.23 1.72 0.23 0.17 0.12 GLF Cut Typical Min. C 8.10 6.27 7.61 4.99 1.33 0.64 0.71 0.46 Cut Typical Plough 9.82 10.51 8.41 4.93 1.89 1.61 1.44 0.47 Cut Rested Min. C 7.51 5.65 6.91 4.76 1.13 0.37 0.35 0.61 Graz. Typical Min. C 9.36 6.26 6.91 5.2 0.37 0.28 0.63 0.22 Graz. Typical Plough 9.27 8.08 7.31 4.85 0.58 0.64 0.77 0.39 Graz. Rested Min. C 15.90 5.75 6.2 4.58 1.15 0.29 0.32 0.14

Herbage magnesium (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 1.59 1.48 1.65 1.21 0.11 0.08 0.08 0.08 Graz. Typical n/a 1.98 1.58 1.66 1.33 0.08 0.07 0.04 0.08 Grass Cut Typical Min. C 1.22 1.29 1.43 1.13 0.07 0.10 0.06 0.05 Cut Typical Plough 0.93 1.17 1.23 0.8 0.05 0.11 0.07 0.07 Cut Rested Min. C 1.43 1.17 1.29 1.16 1.00 0.05 0.04 0.09 Graz. Typical Min. C 1.97 1.42 1.55 1.39 0.03 0.07 0.05 0.07 Graz. Typical Plough 1.44 1.23 1.29 1.2 0.09 0.09 0.07 0.04 Graz. Rested Min. C 2.04 1.38 1.49 1.41 0.05 0.08 0.06 0.09 GL Cut Typical Min. C 1.76 1.32 1.56 1.06 0.26 0.10 0.13 0.05 Cut Rested Min. C 1.62 1.51 1.35 1.1 0.19 0.16 0.06 0.09 Graz. Typical Min. C 2.24 1.51 1.61 1.36 0.05 0.11 0.09 0.05 Graz. Rested Min. C 2.61 1.39 1.54 1.25 0.18 0.05 0.03 0.05 GLF Cut Typical Min. C 1.81 1.49 1.64 1.31 0.22 0.14 0.11 0.13 Cut Typical Plough 1.84 2.12 1.89 1.21 0.28 0.20 0.20 0.14 Cut Rested Min. C 1.91 1.35 1.51 1.262 0.26 0.07 0.06 0.15 Graz. Typical Min. C 2.22 1.45 1.6 1.31 0.11 0.04 0.07 0.05 Graz. Typical Plough 2.08 1.77 1.61 1.28 0.10 0.09 0.12 0.06 Graz. Rested Min. C 2.94 1.52 1.59 1.31 0.12 0.07 0.05 0.04

159

Herbage potassium (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 18.57 17.04 14.31 11.89 1.13 1.58 1.05 1.44 Graz. Typical n/a 23.83 17.48 17.51 19.86 1.52 1.48 0.73 1.32 Grass Cut Typical Min. C 16.51 16.62 14.04 10.14 0.46 1.00 0.75 0.81 Cut Typical Plough 17.13 18.07 18.56 12.77 0.84 0.49 1.02 1.39 Cut Rested Min. C 17.46 16.07 13.65 12.99 0.64 0.88 0.52 1.54 Graz. Typical Min. C 24.94 18.24 19.1 21.86 0.87 1.34 0.55 0.99 Graz. Typical Plough 24.74 18.94 20.12 21.26 1.48 0.85 1.20 1.29 Graz. Rested Min. C 22.17 19.14 17.52 23.06 1.02 0.79 0.68 1.46 GL Cut Typical Min. C 17.77 17.04 16.51 12.93 1.11 0.75 1.05 1.57 Cut Rested Min. C 18.31 17.54 15.1 14.21 0.98 1.31 0.68 1.80 Graz. Typical Min. C 25.35 18.39 18.56 22.8 0.89 0.90 0.73 1.34 Graz. Rested Min. C 22.05 20.14 19.3 22.26 3.12 1.19 0.76 1.09 GLF Cut Typical Min. C 17.57 16.39 15.71 13.74 1.18 0.63 0.64 1.45 Cut Typical Plough 20.85 17.43 17.7 14.07 1.64 1.22 0.82 1.93 Cut Rested Min. C 15.83 16.42 13.23 12.66 1.34 0.98 0.72 1.22 Graz. Typical Min. C 23.77 18.04 18.14 21.39 2.13 1.22 1.43 0.85 Graz. Typical Plough 24.7 22.7 21.19 22.65 2.80 1.37 1.18 1.07 Graz. Rested Min. C 17.03 18.69 17.46 21.29 1.63 0.97 0.55 0.78

Herbage sodium (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 1.49 2.56 2.84 1.89 0.28 0.62 0.44 0.23 Graz. Typical n/a 1.09 1.51 1.79 0.9 0.21 0.25 0.25 0.09 Grass Cut Typical Min. C 1.19 1.43 2.0 1.6 0.21 0.25 0.35 0.21 Cut Typical Plough 0.83 1.04 1.33 0.89 0.13 0.12 0.11 0.10 Cut Rested Min. C 1.49 0.93 1.71 1.4 0.32 0.11 0.30 0.24 Graz. Typical Min. C 1.05 0.90 1.2 0.73 0.14 0.12 0.19 0.04 Graz. Typical Plough 0.61 0.60 1.1 0.83 0.05 0.04 0.11 0.08 Graz. Rested Min. C 1.56 0.78 1.13 0.78 0.33 0.08 0.13 0.04 GL Cut Typical Min. C 1.24 1.28 2.0 1.01 0.23 0.24 0.35 0.12 Cut Rested Min. C 1.31 1.23 1.38 1.14 0.31 0.26 0.16 0.19 Graz. Typical Min. C 1.35 1.21 1.21 0.71 0.26 0.24 0.14 0.04 Graz. Rested Min. C 1.71 0.88 1.13 0.78 0.30 0.19 0.13 0.09 GLF Cut Typical Min. C 1.059 1.83 2.05 2.11 0.11 0.34 0.32 0.33 Cut Typical Plough 1.07 2.24 2.39 1.7 0.12 0.26 0.33 0.22 Cut Rested Min. C 1.23 1.72 2.99 1.98 0.10 0.20 0.26 0.40 Graz. Typical Min. C 1.79 1.30 1.85 0.98 0.38 0.27 0.29 0.12 Graz. Typical Plough 1.42 1.78 2.24 1.13 0.10 0.32 0.30 0.18 Graz. Rested Min. C 1.82 1.47 1.78 0.89 0.26 0.21 0.18 0.03

160

Herbage DOMD (g kg-1 DM)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 607.6 611.3 615.2 551.7 10.44 9.27 11.4 16.44 Graz. Typical n/a 559.5 615.5 638.5 591.6 7.71 13.78 7.13 7.77 Grass Cut Typical Min. C 610.2 614.3 610.8 566.6 8.90 8.01 15.3 14.4 Cut Typical Plough 609.7 638.1 653.5 565.4 13.57 7.16 5.00 14.4 Cut Rested Min. C 594.6 609.9 631.9 577.0 10.05 13.72 10.8 15.1 Graz. Typical Min. C 614.3 608.2 641.2 574.4 7.50 11.79 5.76 11.0 Graz. Typical Plough 635.9 622.8 660.8 605.3 5.98 13.15 6.44 10.2 Graz. Rested Min. C 600.6 614.0 624.6 600.3 11.77 10.17 10.5 13.2 GL Cut Typical Min. C 609.5 613.7 634.1 578.2 6.46 7.82 4.45 17.6 Cut Rested Min. C 601.3 600.7 629.1 571.5 6.44 13.36 11.6 15.7 Graz. Typical Min. C 646.8 627.0 649.7 596.7 7.17 15.24 9.08 9.4 Graz. Rested Min. C 664.9 611.7 622.1 582.1 5.03 12.3 10.8 12.2 GLF Cut Typical Min. C 608.7 608.8 630.0 567.3 6.28 16.44 9.20 21.2 Cut Typical Plough 610.9 606.9 631.4 571.1 7.66 5.52 5.52 18.6 Cut Rested Min. C 601.6 599.5 621.1 565.7 6.24 11.02 12.2 19.5 Graz. Typical Min. C 649.1 610.2 652.0 601.7 4.21 12.03 6.38 12.73 Graz. Typical Plough 656.1 646.9 668.5 583.3 6.08 8.05 5.86 14.9 Graz. Rested Min. C 657.2 614.3 629.0 594.7 11.52 7.67 8.03 10.9

161

Appendix 1f: North Wyke summed percentage cover of grasses, legumes and forbs.

Sown grass species summed percentage cover (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 68.1 68.7 50.8 15.4 12.7 7.4 6.7 4.3 Grassland Graz. Typical n/a 36.2 50.7 26.9 10.2 6.5 7.2 4.0 4.4 Grass Cut Typical Min. C 77.1 67.7 51.8 53.7 8.4 5.6 9.6 8.6 Cut Typical Plough 85.5 97.8 87.1 86.0 8.5 4.5 3.8 6.2 Cut Rested Min. C 54.9 65.1 57.5 62.5 13.1 6.1 4.5 9.8 Graz. Typical Min. C 37.8 54.3 23.7 14.1 10.1 6.6 6.3 3.7 Graz. Typical Plough 87.7 94.0 88.6 61.5 2.5 4.3 2.8 9.9 Graz. Rested Min. C 63.0 61.3 32.7 21.8 11.8 4.0 8.7 4.5 GL Cut Typical Min. C 79.0 75.0 77.5 57.8 8.3 4.8 1.3 9.2 Cut Rested Min. C 80.4 66.5 50.0 54.3 9.8 9.9 4.8 7.6 Graz. Typical Min. C 45.2 57.7 29.3 22.7 12.7 5.0 4.5 8.7 Graz. Rested Min. C 62.4 73.5 48.5 24.1 12.6 8.7 6.8 8.1 GLF Cut Typical Min. C 65.9 74.3 65.0 42.3 11.7 7.1 3.3 5.4 Cut Typical Plough 67.3 59.8 77.6 78.5 7.8 6.8 4.5 4.7 Cut Rested Min. C 66.3 72.5 61.3 48.7 10.5 9.6 2.9 7.7 Graz. Typical Min. C 38.2 64.6 32.1 20.7 12.8 4.4 7.3 8.3 Graz. Typical Plough 75.6 55.0 70.5 42.0 10.8 5.7 4.0 4.9 Graz. Rested Min. C 74.0 53.3 24.5 23.7 5.67 5.2 3.0 10.3

Sown non-legume forb species summed percentage cover (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0 0.8 0 0 0 0.8 0 Grassland Graz. Typical n/a 0.1 0 0.1 0 0.1 0 0.1 0 Grass Cut Typical Min. C 0 0 0 0 0 0 0 0 Cut Typical Plough 0 0 0 0 0 0 0 0 Cut Rested Min. C 0 0 0 0.01 0 0 0 0.01 Graz. Typical Min. C 0 0 0 0.1 0 0 0 0.1 Graz. Typical Plough 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0 0 0 0 0 0 0 0 GL Cut Typical Min. C 0 0.1 0 0 0 0.1 0 0 Cut Rested Min. C 0 0.1 0 0.1 0 0.1 0 0.1 Graz. Typical Min. C 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0.2 0.2 0.1 0 0.2 0.2 0.1 0 GLF Cut Typical Min. C 3.7 6.5 3.2 3.0 1.8 3.4 1.4 2.3 Cut Typical Plough 39.8 44.3 24.8 12.1 3.1 7.8 4.5 3.0 Cut Rested Min. C 4.8 2.5 3.0 1.2 2.5 1.2 1.2 0.4 Graz. Typical Min. C 3.9 5.0 5.6 1.9 1.3 1.3 2.8 1.3 Graz. Typical Plough 15.3 22.4 10.1 1.8 2.4 4.5 1.9 1.2 Graz. Rested Min. C 6.3 6.1 3.9 0.9 2.5 1.7 1.4 0.6

162

Sown legume species summed percentage cover (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 31.2 5.5 1.1 0 15.3 1.8 0.6 0 Grassland Graz. Typical n/a 15.8 2.8 7.0 0.4 7.3 1.6 5.4 0.1 Grass Cut Typical Min. C 0 0.03 0 0 0 0.03 0 0 Cut Typical Plough 0 0 0.2 2.8 0 0 0.1 1.6 Cut Rested Min. C 0.1 0 0 0 0.1 0 0 0 Graz. Typical Min. C 0 0.1 0.03 0 0 0.1 0.03 0 Graz. Typical Plough 0 0 0.5 3.6 0 0 0.4 1.3 Graz. Rested Min. C 0 0.03 0 0 0 0.03 0 0 GL Cut Typical Min. C 43.0 15.2 9.8 0.01 12.4 5.3 6.2 0.01 Cut Rested Min. C 51.0 8.9 2.8 0.01 17.3 2.2 0.4 0.01 Graz. Typical Min. C 28.5 10.4 5.8 0.7 12.4 2.4 2.0 0.3 Graz. Rested Min. C 40.3 4.0 4.7 0.03 13.3 1.0 2.7 0.03 GLF Cut Typical Min. C 40.9 15.8 2.2 0.03 10.1 3.0 0.8 0.03 Cut Typical Plough 62.4 27.1 11.8 1.4 11.4 6.2 4.6 0.8 Cut Rested Min. C 54.0 3.2 1.3 0 7.3 1.0 0.4 0 Graz. Typical Min. C 30.9 10.6 11.1 0.1 4.5 2.5 3.6 0.1 Graz. Typical Plough 25.4 39.6 36.9 0.2 4.0 7.9 4.1 0.2 Graz. Rested Min. C 45.7 3.3 1.6 0.1 7.8 1.5 0.8 0.1

163

Appendix 1g: North Wyke species richness of grasses, legumes and non-legume forbs.

Sown grass species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 2.5 1.8 1.3 1.3 0.3 0.5 0.3 0.3 Grassland Graz. Typical n/a 2 1.8 1.3 1.5 0.7 0.3 0.3 0.3 Grass Cut Typical Min. C 3.5 3.5 3.3 4 0.3 0.3 0.3 0 Cut Typical Plough 4 3.5 3.5 5 0.4 0.3 0.3 0 Cut Rested Min. C 1.8 3.3 3 4 0.6 0.3 0 0.4 Graz. Typical Min. C 3.3 3.8 3.3 3 0.3 0.3 0.3 0 Graz. Typical Plough 3.5 3.3 3.5 4.5 0.6 0.3 0.3 0.3 Graz. Rested Min. C 3.3 3.3 3.3 3 0.5 0.6 0.3 0 GL Cut Typical Min. C 2.3 3.5 3.5 4 0.5 0.3 0.3 0 Cut Rested Min. C 2.5 3.3 3.5 3.3 0.3 0.3 0.3 0.3 Graz. Typical Min. C 3.3 2.3 3.3 3 0.5 0.3 0.3 0 Graz. Rested Min. C 3.3 2.8 3.8 3.3 0.3 0.3 0.5 0.3 GLF Cut Typical Min. C 2.5 3.5 3.3 3.5 0.3 0.5 0.3 0.3 Cut Typical Plough 4.3 3.3 3.3 4.8 0.5 0.3 0.3 0.3 Cut Rested Min. C 3.3 3.5 3.3 3.8 0.5 0.3 0.3 0.3 Graz. Typical Min. C 3.8 3 3 3 0.3 0 0 0 Graz. Typical Plough 3.8 3 3.5 4.3 0.3 0 0.5 0.5 Graz. Rested Min. C 3 3 3.3 2.3 0.4 0.4 0.3 0.3

Sown non-legume forb species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0 03 0 0 0 0.3 0 Grassland Graz. Typical n/a 0.3 0 0.3 0 0.3 0 0.3 0 Grass Cut Typical Min. C 0 0 0 0 0 0 0 0 Cut Typical Plough 0 0 0 0 0 0 0 0 Cut Rested Min. C 0 0 0 0.3 0 0 0 0.3 Graz. Typical Min. C 0 0 0 0.3 0 0 0 0.3 Graz. Typical Plough 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0 0 0 0 0 0 0 0 GL Cut Typical Min. C 0 0.3 0 0 0 0.3 0 0 Cut Rested Min. C 0 0.3 0 0.3 0 0.3 0 0.3 Graz. Typical Min. C 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0.8 0.3 0.3 0 0.8 0.3 0.3 0 GLF Cut Typical Min. C 4.5 3.8 3 2.3 1.5 1.3 1.0 0.9 Cut Typical Plough 6 5.8 5.5 4 0 0.3 0.3 0.4 Cut Rested Min. C 4 2.3 2.5 1.5 1.4 0.9 1.0 0.5 Graz. Typical Min. C 5.3 4.3 3.8 1.3 0.8 0.5 0.6 0.8 Graz. Typical Plough 6 5.8 5.3 1.3 0 0.3 0.3 0.6 Graz. Rested Min. C 5.5 3.8 3.8 1.5 0.5 0.6 1.0 0.6

164

. Sown legume species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 1.0 1.0 0.8 0 0 0 0.3 0 Grassland Graz. Typical n/a 0.8 1.0 1.0 1.0 0.3 0 0 0 Grass Cut Typical Min. C 0 0.3 0 0 0 0.3 0 0 Cut Typical Plough 0 0 0.8 0.8 0 0 0.3 0.3 Cut Rested Min. C 0.3 0 0 0 0.3 0 0 0 Graz. Typical Min. C 0 0.3 0.3 0 0 0.3 0.3 0 Graz. Typical Plough 0 0 0.8 1.3 0 0 0.5 0.3 Graz. Rested Min. C 0 0.3 0 0 0 0.3 0 0 GL Cut Typical Min. C 3.8 2.3 1.5 0.5 0.3 0.3 0.5 0.3 Cut Rested Min. C 2.8 2.5 2.0 0.3 0.5 0.3 0 0.3 Graz. Typical Min. C 3.0 1.8 1.3 0.8 0.4 0.8 0.3 0.3 Graz. Rested Min. C 2.5 1.5 1.0 0.3 0.5 0.3 0 0.3 GLF Cut Typical Min. C 3.3 2.3 2.0 0.5 0.5 0.3 0 0.3 Cut Typical Plough 4.0 4.0 2.8 1.5 0 0 0.3 0.3 Cut Rested Min. C 3.3 2.0 1.3 0 0.6 0 0.5 0 Graz. Typical Min. C 3.5 2.3 1.8 0.5 0.3 0.3 0.3 0.3 Graz. Typical Plough 4.3 3.8 2.5 0.5 0.3 0.3 0.3 0.5 Graz. Rested Min. C 3.3 1.8 0.8 0.5 0.3 0.3 0.3 0.3

165

Appendix 1h: North Wyke density of legumes and non-legume forb flower heads

Non-legume forb flower density (m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0.8 0 0 0 0.6 0 0 Grassland Graz. Typical n/a 1.2 5.2 0 0 0.8 2.0 0 0 Grass Cut Typical Min. C 0.7 1.8 0 0 0.4 1.2 0 0 Cut Typical Plough 0.2 5.3 0 0 0.2 1.5 0 0 Cut Rested Min. C 0 1.2 0 0 0 0.7 0 0 Graz. Typical Min. C 0.2 3.7 0 0 0.2 1.3 0 0 Graz. Typical Plough 0.2 15.3 0 0.5 0.2 1.7 0 0.5 Graz. Rested Min. C 0.2 1.8 0 0 0.2 1.1 0 0 GL Cut Typical Min. C 0 0.3 0 0 0 0.2 0 0 Cut Rested Min. C 0.5 1.2 0.2 0 0.5 1.0 0.2 0 Graz. Typical Min. C 0.3 2.0 0 0 0.2 0.7 0 0 Graz. Rested Min. C 0.2 6.7 0.2 0 0.2 3.8 0.2 0 GLF Cut Typical Min. C 0 14.2 2.8 5.0 0 7.7 2.2 2.9 Cut Typical Plough 3.3 26.8 6.2 11.2 1.4 7.2 2.7 2.5 Cut Rested Min. C 1.2 11.5 2.8 4.2 0.8 6.7 1.9 4.2 Graz. Typical Min. C 0.3 14.8 7.5 2.5 0.2 4.9 4.7 2.5 Graz. Typical Plough 2.7 22.3 8.5 2.8 1.2 3.5 4.9 2.8 Graz. Rested Min. C 0.2 24.8 11.5 8.8 0.2 8.1 7.5 5.2

Legume forb flower density (m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 16.0 25.0 1.2 0 6.7 9.2 0.7 0 Grassland Graz. Typical n/a 13.7 0.7 4.3 0 3.0 0.7 3.3 0 Grass Cut Typical Min. C 3.8 0 0 0 3.8 0 0 0 Cut Typical Plough 0 0 1.8 0 0 0 1.3 0 Cut Rested Min. C 0 0 0 0 0 0 0 0 Graz. Typical Min. C 0 0 0 0 0 0 0 0 Graz. Typical Plough 0 0.3 7.8 3.0 0 0.3 6.3 3.0 Graz. Rested Min. C 0 0 0 0 0 0 0 0 GL Cut Typical Min. C 58.8 64.8 10.8 0 6.1 18.4 6.8 0 Cut Rested Min. C 52.7 20.8 5.3 0 13.1 4.4 1.4 0 Graz. Typical Min. C 26.3 13.7 10.5 0 6.3 4.5 3.5 0 Graz. Rested Min. C 52.0 5.2 12.3 0 13.0 1.6 10.8 0 GLF Cut Typical Min. C 85.8 37.3 1.8 0 22.5 9.3 0.7 0 Cut Typical Plough 145.0 92.3 10.7 2.3 22.1 20.8 2.8 2.1 Cut Rested Min. C 79.3 15.3 0.7 0 29.7 4.6 0.7 0 Graz. Typical Min. C 14.7 10.3 5.8 0.2 2.3 4.3 2.5 0.2 Graz. Typical Plough 12.5 58.0 32.8 0.5 5.9 0.9 5.3 0.5 Graz. Rested Min. C 65.0 5.7 2.2 0 6.6 2.3 2.0 0

166

Appendix 1i: North Wyke density of grasses, legumes and non-legume forb winter seed heads.

Winter grass seed head density (seed heads m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0.5 1.3 0 3.5 0.3 0.7 0 2.1 Grassland Graz. Typical n/a 3.3 2.3 13.2 5.1 1.2 0.6 5.6 3.8 Grass Cut Typical Min. C 0.7 1.2 0 0.3 0 1.2 0 0.2 Cut Typical Plough 2.0 0.2 0 0.3 0.7 0.2 0 0.2 Cut Rested Min. C 0 0.2 0 1.7 0 0.2 0 0.8 Graz. Typical Min. C 5.0 2.1 15.5 2.5 0.4 1.7 6.9 1.4 Graz. Typical Plough 9.2 5.1 12.1 6.9 2.3 1.4 2.4 2.5 Graz. Rested Min. C 26.1 13.4 24.4 2.0 2.7 3.8 7.9 0.5 GL Cut Typical Min. C 1.2 0.5 0 3.1 0.8 0.5 0 1.6 Cut Rested Min. C 0.3 0.8 0 3.3 0.3 0.5 0 1.3 Graz. Typical Min. C 11.2 4.5 7.8 1.8 1.2 2.6 1.1 0.4 Graz. Rested Min. C 25.2 7.9 16.7 5.1 2.9 2.3 5.3 1.2 GLF Cut Typical Min. C 0 0.17 0 4.0 0 0.17 0 1.3 Cut Typical Plough 3.0 0 0 6.6 1.7 0 0 2.3 Cut Rested Min. C 0 0 0 1.7 0 0 0 1.0 Graz. Typical Min. C 10.6 2.8 10.7 1.5 5.1 2.4 4.5 0.5 Graz. Typical Plough 10.4 1.3 9.9 4.1 2.5 0.6 4.0 1.9 Graz. Rested Min. C 11.9 2.5 10.1 2.8 5.1 1.0 1.8 1.3

Winter non-legume forb seed head density (seed heads m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0.2 0 0 0 0.2 0 0 0 Grassland Graz. Typical n/a 0.2 0 0 0 0.2 0 0 0 Grass Cut Typical Min. C 1.5 0.2 0 0 1.5 0.2 0 0 Cut Typical Plough 3.0 0 0 0 2.8 0 0 0 Cut Rested Min. C 0 0 0 0 0 0 0 0 Graz. Typical Min. C 11.7 0.3 0 0 3.6 0.3 0 0 Graz. Typical Plough 2.3 1.8 0 0 1.3 0.3 0 0 Graz. Rested Min. C 0.8 0 0 0 0.8 0 0 0 GL Cut Typical Min. C 4.0 0 0 0 2.8 0 0 0 Cut Rested Min. C 0 0 0 0 0 0 0 0 Graz. Typical Min. C 3.1 0.2 0 0 2.3 0.2 0 0 Graz. Rested Min. C 0 0 0 0 0 0 0 0 GLF Cut Typical Min. C 4.5 1.7 0 0 4.5 1.1 0 Cut Typical Plough 9.2 3.6 0 2.3 6.7 1.3 0 1.7 Cut Rested Min. C 0 0.8 0 0 0 0.8 0 Graz. Typical Min. C 22.3 3.6 0.7 0.7 13.5 2.5 0.5 0.7 Graz. Typical Plough 45.5 10.7 10.7 2.5 11.1 4.0 5.8 2.5 Graz. Rested Min. C 36.0 21.1 12.7 3.1 18.5 10.8 10.7 1.9

167

Winter legume seed head density (seed heads m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0 0 0 0 0 0 0 Grassland Graz. Typical n/a 2.3 0 0 0 1.6 0 0 0 Grass Cut Typical Min. C 0 0 0 0 0 0 0 0 Cut Typical Plough 0 0 0 0 0 0 0 0 Cut Rested Min. C 0 0 0 0 0 0 0 0 Graz. Typical Min. C 0 0 0 0 0 0 0 0 Graz. Typical Plough 3.6 0 0 0 3.6 0 0 0 Graz. Rested Min. C 0 0 0 0 0 0 0 0 GL Cut Typical Min. C 3.0 0.2 0 0 1.2 0.2 0 0 Cut Rested Min. C 0.3 0 0 0 0.3 0 0 0 Graz. Typical Min. C 5.0 0.7 0 0 1.5 0.7 0 0 Graz. Rested Min. C 14.5 0 0 0 6.2 0 0 0 GLF Cut Typical Min. C 4.8 0 0 0 1.5 0 0 0 Cut Typical Plough 8.1 0.8 0 0 2.8 0.6 0 0 Cut Rested Min. C 0.2 0 0 0 0.2 0 0 0 Graz. Typical Min. C 4.1 0.2 0 0 1.7 0.2 0 0 Graz. Typical Plough 8.4 0.5 0 0 2.1 0.2 0 0 Graz. Rested Min. C 27.9 0 0 0 2.7 0 0 0

168

Appendix 1j: North Wyke beetle biomass (mg)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 5.9 67.3 21.7 15.6 2.0 12.8 9.9 4.9 Grassland Graz. Typical n/a 21.0 27.7 8.1 2.9 8.7 10.4 3.3 1.5 Grass Cut Typical Min. C 1.6 47.9 11.7 4.9 0.6 10.2 4.0 2.6 Cut Typical Plough 5.9 28.3 21.7 7.0 1.9 5.9 4.9 3.2 Cut Rested Min. C 14.4 31.2 22.3 5.8 3.0 3.48 2.5 1.9 Graz. Typical Min. C 3.7 15.6 10.9 16.6 1.2 4.0 3.6 8.3 Graz. Typical Plough 8.9 27.8 6.7 7.6 3.5 6.7 1.8 4.2 Graz. Rested Min. C 34.8 68.4 12.1 3.3 6.7 31.5 2.5 0.5 GL Cut Typical Min. C 6.3 98.3 39.4 14.2 2.0 30.9 4.7 5.5 Cut Rested Min. C 48.4 37.7 20.6 11.3 24.5 7.2 7.4 3.6 Graz. Typical Min. C 41.2 57.3 7.1 5.0 2.1 11.6 1.9 2.7 Graz. Rested Min. C 31.5 60.8 38.2 9.8 7.4 7.06 24.9 3.9 GLF Cut Typical Min. C 9.7 64.9 17.6 4.6 6.6 15.0 6.3 1.4 Cut Typical Plough 4.4 98.6 33.5 10.7 0.9 13.3 6.0 3.2 Cut Rested Min. C 49.4 56.1 19.8 10.9 23.4 15.9 6.1 4.4 Graz. Typical Min. C 8.3 36.3 14.6 4.3 1.0 6.6 3.9 2.4 Graz. Typical Plough 23.4 110.5 33.7 11.9 4.3 41.3 8.4 3.9 Graz. Rested Min. C 31.1 54.5 12.2 4.0 6.9 10.4 2.4 2.1

169

Appendix 1k: North Wyke beetle and pollinator abundance

Beetle abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 6.3 48.8 11.3 7.5 2.7 8.3 3.1 1.8 Grassland Graz. Typical n/a 19.5 41.0 10.0 10.3 6.6 13.9 1.7 5.1 Grass Cut Typical Min. C 2.5 35.5 8.0 5.8 0.6 10.7 2.2 2.3 Cut Typical Plough 2.8 25.3 15.8 13.0 1.0 4.9 4.3 3.7 Cut Rested Min. C 19.0 25.8 9.0 3.8 3.6 7.2 1.0 1.2 Graz. Typical Min. C 6.5 21.8 8.3 8.0 1.3 5.0 1.4 1.8 Graz. Typical Plough 6.3 21.3 13.0 11.0 2.2 1.8 1.9 2.3 Graz. Rested Min. C 14.0 40.8 10.5 7.3 1.1 6.5 3.0 1.8 GL Cut Typical Min. C 8.0 76.5 23.5 12.8 1.8 11.6 4.2 1.9 Cut Rested Min. C 25.5 55.5 16.5 9.8 4.2 11.8 5.6 2.1 Graz. Typical Min. C 24.0 50.3 13.3 6.0 4.5 8.7 2.4 2.2 Graz. Rested Min. C 27.0 33.5 12.0 7.0 6.5 5.2 3.4 2.3 GLF Cut Typical Min. C 10.0 62.0 16.8 8.0 7.4 19.8 2.9 2.0 Cut Typical Plough 4.0 70.5 21.8 14.8 0.7 5.3 1.0 1.8 Cut Rested Min. C 23.0 80.8 11.5 7.0 3.9 29.0 3.4 2.0 Graz. Typical Min. C 11.3 39.0 14.5 6.5 0.9 5.8 0.6 1.8 Graz. Typical Plough 19.0 68.3 34.0 19.5 3.4 16.7 9.3 3.5 Graz. Rested Min. C 27.0 44.5 13.3 8.5 7.5 8.9 3.4 2.0

Total pollinator abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 2.0 3.3 0.8 1.5 0.8 1.7 0.8 1.5 Grassland Graz. Typical n/a 2.5 0.3 0 0.3 1.3 0.3 0 0.3 Grass Cut Typical Min. C 0.8 0.5 6.0 8.5 0.5 0.5 4.5 5.9 Cut Typical Plough 1.3 2.0 0.5 3.8 1.3 1.1 0.5 3.8 Cut Rested Min. C 3.3 0.5 0.3 0 3.0 0.5 0.3 0 Graz. Typical Min. C 0.8 0.5 4.5 0 0.5 0.5 3.9 0 Graz. Typical Plough 0.3 1.0 1.0 0 0.3 0.7 0.7 0 Graz. Rested Min. C 3.0 1.0 2.3 0 2.7 0.6 1.9 0 GL Cut Typical Min. C 14.8 12.0 1.0 4.3 3.2 4.5 0.7 4.0 Cut Rested Min. C 17.8 3.0 1.8 0 8.1 1.1 1.0 0 Graz. Typical Min. C 4.0 1.8 0 4.3 1.2 0.6 0 4.3 Graz. Rested Min. C 12.0 1.3 1.5 2.8 5.9 0.6 1.0 2.8 GLF Cut Typical Min. C 15.8 14.0 6.5 0 6.6 3.1 6.2 0 Cut Typical Plough 45.0 73.3 2.0 0 9.1 10.5 1.7 0 Cut Rested Min. C 16.8 4.5 3.8 0 3.0 1.8 2.3 0 Graz. Typical Min. C 4.3 4.3 0.5 1.3 1.1 1.7 0.5 1.3 Graz. Typical Plough 4.8 12.5 0 1.3 1.1 1.2 0 1.3 Graz. Rested Min. C 17.8 16.5 2.3 2.3 4.7 9.2 1.9 2.3

170

Bumblebee abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 2.0 1.8 0.3 0 0.8 1.4 0.3 0 Grassland Graz. Typical n/a 1.8 0 0 0 1.2 0 0 0 Grass Cut Typical Min. C 0.3 0 1.3 0 0.3 0 0.8 0 Cut Typical Plough 1.0 0 0.3 0.3 1.0 0 0.3 0.3 Cut Rested Min. C 2.8 0 0 0 2.8 0 0 0 Graz. Typical Min. C 0.3 0 0.8 0 0.3 0 0.8 0 Graz. Typical Plough 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0.3 0 0.3 0 0.3 0 0.3 0 GL Cut Typical Min. C 10.3 10.0 0 0 2.9 4.1 0 0 Cut Rested Min. C 12.8 2.0 0 0 5.5 0.8 0 0 Graz. Typical Min. C 3.3 1.3 0 0 1.0 0.5 0 0 Graz. Rested Min. C 6.8 0.8 0 0 3.0 0.5 0 0 GLF Cut Typical Min. C 12.5 7.8 1.3 0 5.3 1.1 1.3 0 Cut Typical Plough 23.0 53.8 0.5 0 5.8 9.5 0.3 0 Cut Rested Min. C 12.8 2.3 0.8 0 2.4 1.1 0.8 0 Graz. Typical Min. C 3.5 2.0 0 0 0.9 1.4 0 0 Graz. Typical Plough 3.3 8.3 0 0.3 1.5 1.3 0 0.3 Graz. Rested Min. C 12.0 7.3 0 0 2.6 4.9 0 0

Butterfly abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0.3 0.3 0 0 0.3 0.3 0 Grassland Graz. Typical n/a 0.8 0.3 0 0 0.5 0.3 0 0 Grass Cut Typical Min. C 0.5 0.3 0.5 2.3 0.5 0.3 1.2 1.3 Cut Typical Plough 0 0.3 0.3 1.8 0 0.3 0.3 1.8 Cut Rested Min. C 0.5 0.5 0.3 0 0.3 0.5 0.3 0 Graz. Typical Min. C 0.5 0.5 0.8 0 0.3 0.5 0.5 0 Graz. Typical Plough 0.3 0 1.0 0 0.3 0 0.7 0 Graz. Rested Min. C 0.8 0.5 0.5 0 0.5 0.5 0.5 0 GL Cut Typical Min. C 2.5 0.3 0.5 0.3 1.6 0.5 0.3 0.3 Cut Rested Min. C 1.8 0.8 1.8 0 1.1 0.5 1.0 0 Graz. Typical Min. C 0 0.5 0 0.5 0 0.3 0 0.5 Graz. Rested Min. C 1.0 0 0.8 0.3 0.7 0 0.5 0.3 GLF Cut Typical Min. C 0.8 2.0 3.5 0 0.8 0.7 3.2 0 Cut Typical Plough 6.0 4.0 0.8 0 1.8 2.0 0.8 0 Cut Rested Min. C 2.3 0.8 2.3 0 0.3 0.3 1.4 0 Graz. Typical Min. C 0.3 0.3 0.5 0 0.3 0.3 0.5 0 Graz. Typical Plough 0.3 0.3 0 0 0.3 0.3 0 0 Graz. Rested Min. C 1.8 1.0 1.0 0.5 0.8 0.6 0.7 0.5

171

Honeybee abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0.5 0.3 0 0 0.5 0.3 0 Grassland Graz. Typical n/a 0 0 0 0 0 0 0 0 Grass Cut Typical Min. C 0 0 0 0 0 0 0 0 Cut Typical Plough 0 0 0 0 0 0 0 0 Cut Rested Min. C 0 0 0 0 0 0 0 0 Graz. Typical Min. C 0 0 0.5 0 0 0 0.5 0 Graz. Typical Plough 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0 0 0 0 0 0 0 0 GL Cut Typical Min. C 1.8 0.3 0 3.5 0.5 0.3 0 3.5 Cut Rested Min. C 3.3 0 0 0 2.6 0 0 0 Graz. Typical Min. C 0.8 0 0 2.8 0.5 0 0 2.8 Graz. Rested Min. C 0.8 0 0.3 2.3 2.3 0 0.3 2.3 GLF Cut Typical Min. C 2.0 2.0 0 0 0.7 0.8 0 0 Cut Typical Plough 12.8 7.8 0 0 3.8 4.3 0 0 Cut Rested Min. C 1.3 0.5 0 0 0.8 0.3 0 0 Graz. Typical Min. C 0.3 0.5 0 0 0.3 0.5 0 0 Graz. Typical Plough 1.3 0 0 0.8 0.9 0 0 0.8 Graz. Rested Min. C 3.8 3.0 0.3 1.3 1.5 2.4 0.3 1.3

Hoverfly abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0.8 0 1.5 0 0.8 0 1.5 Grassland Graz. Typical n/a 0 0 0 0.3 0 0 0 0.3 Grass Cut Typical Min. C 0 0.3 3.3 6.3 0 0.3 2.9 5.0 Cut Typical Plough 0.3 1.8 0 1.8 0.3 1.2 0 1.8 Cut Rested Min. C 0 0 0 0 0 0 0 0 Graz. Typical Min. C 0 0 1.5 0 0 0 1.5 0 Graz. Typical Plough 0 1.0 0 0 0 0.7 0 0 Graz. Rested Min. C 0 0.5 1.5 0 0 0.5 1.5 0 GL Cut Typical Min. C 0.3 0.5 0.5 0.5 0.3 0.3 0.5 0.3 Cut Rested Min. C 0 0.3 0 0 0 0.3 0 0 Graz. Typical Min. C 0 0 0 1.0 0 0 0 1.0 Graz. Rested Min. C 0.5 0.5 0.5 0.3 0.3 0.3 0.3 0.3 GLF Cut Typical Min. C 0.5 2.3 1.8 0 0.3 0.9 1.8 0 Cut Typical Plough 3.3 7.8 0.8 0 1.9 2.6 0.8 0 Cut Rested Min. C 0.5 1.0 0.8 0 0.5 0.4 0.8 0 Graz. Typical Min. C 0.3 1.5 0 1.3 0.3 1.0 0 1.3 Graz. Typical Plough 0 3.8 0 0.3 0 1.9 0 0.3 Graz. Rested Min. C 0.3 5.3 1.0 0.5 0.3 3.5 1.0 0.5

172

Appendix 1l: North Wyke beetle and pollinator species richness.

Beetle species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 3.8 7.0 6.8 4.0 1.2 1.5 1.8 0.4 Grassland Graz. Typical n/a 6.5 7.8 5.3 2.3 1.3 1.1 0.9 0.6 Grass Cut Typical Min. C 1.5 9.8 4.5 2.3 0.3 1.3 1.0 0.8 Cut Typical Plough 2.3 8.8 6.8 3.3 0.8 1.9 1.3 0.8 Cut Rested Min. C 7.5 8.0 5.8 3.0 1.0 0.6 0.8 0.7 Graz. Typical Min. C 3.8 5.5 4.5 3.5 0.6 0.6 0.6 0.3 Graz. Typical Plough 3.8 9.8 5.3 4.3 0.9 1.6 0.9 0.6 Graz. Rested Min. C 8.8 10.0 5.5 0.3 0.5 0.7 0.3 0.5 GL Cut Typical Min. C 4.8 10.3 9.8 5.0 0.9 1.0 1.7 0.7 Cut Rested Min. C 9.5 10.3 7.5 5.3 1.7 1.1 2.5 1.1 Graz. Typical Min. C 8.5 9.8 4.5 3.0 1.3 0.9 1.0 0.6 Graz. Rested Min. C 8.8 13.8 6.8 4.0 0.8 2.0 1.3 0.7 GLF Cut Typical Min. C 3.5 11.3 8.3 3.8 1.7 0.9 1.4 0.8 Cut Typical Plough 3.5 16.8 9.5 5.8 0.5 1.8 0.6 1.4 Cut Rested Min. C 8.3 11.5 6.3 4.0 0.9 1.7 1.9 1.1 Graz. Typical Min. C 4.0 7.3 6.3 2.8 0.4 0.5 0.8 0.5 Graz. Typical Plough 6.8 11.0 11.0 6.3 1.0 1.1 1.9 0.9 Graz. Rested Min. C 8.0 12.8 6.3 3.8 1.8 0.6 1.2 0.6

Bee (Bombus and Apis mellifera) species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 1.0 0 0.3 0.3 0.4 0 0.3 0.3 Grassland Graz. Typical n/a 1.0 0.3 0.5 0 0.7 0.3 0.5 0 Grass Cut Typical Min. C 0.3 0.8 0.3 0 0.3 0.8 0.3 0 Cut Typical Plough 0.3 0.5 0 0 0.3 0.3 0 0 Cut Rested Min. C 0.8 1.8 0.3 0 0.8 0.9 0.3 0 Graz. Typical Min. C 0.3 0 0.3 0 0.3 0 0.3 0 Graz. Typical Plough 0 0 0 0 0 0 0 0 Graz. Rested Min. C 0.8 0.8 0 0 0.8 0.5 0 0 GL Cut Typical Min. C 3.0 0 0.5 0 0.4 0 0.5 0 Cut Rested Min. C 2.3 2.8 0 0 0.6 0.5 0 0 Graz. Typical Min. C 2.0 0 0 0 0.7 0 0 0 Graz. Rested Min. C 3.0 1.5 0 0 0.6 0.6 0 0 GLF Cut Typical Min. C 3.0 3.0 0.5 0 0.7 0.7 0.3 0 Cut Typical Plough 3.5 2.3 0.5 0 0.3 0.3 0.5 0 Cut Rested Min. C 3.5 1.8 0.5 0 0.5 1.0 0.5 0 Graz. Typical Min. C 1.8 2.0 0 0 0.3 0.7 0 0 Graz. Typical Plough 1.8 1.3 0 0 0.9 0.6 0 0 Graz. Rested Min. C 3.3 1.8 0 0.3 0.3 0.3 0 0.3

173

Butterfly species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original Cut Typical n/a 0 0.3 0.3 0 0 0.3 0.3 0 Grassland Graz. Typical n/a 0.8 0.3 0 0 0.5 0.3 0 0 Grass Cut Typical Min. C 0.3 0.3 0.5 0.8 0.3 0.3 0.3 0.5 Cut Typical Plough 0 0.3 0.3 0.3 0 0.3 0.3 0.3 Cut Rested Min. C 0.5 0.5 0.3 0 0.3 0.5 0.3 0 Graz. Typical Min. C 0.5 0.3 0.5 0 0.3 0.3 0.3 0 Graz. Typical Plough 0.3 0 0.5 0 0.3 0 0.3 0 Graz. Rested Min. C 0.8 0.3 0.3 0 0.5 0.3 0.3 0 GL Cut Typical Min. C 1.0 1.0 0.5 0.3 0.4 0.4 0.3 0.3 Cut Rested Min. C 1.5 0.5 1.0 0 0.9 0.3 0.6 0 Graz. Typical Min. C 0 0.5 0 0.3 0 0.3 0 0.3 Graz. Rested Min. C 1.0 0 0.8 0.3 0.7 0 0.5 0.3 GLF Cut Typical Min. C 0.8 1.3 0.8 0 0.8 0.5 0.5 0 Cut Typical Plough 2.0 2.0 0.3 0 0.4 0.7 0.3 0 Cut Rested Min. C 1.8 0.8 0.8 0 0.3 0.3 0.5 0 Graz. Typical Min. C 0.3 0.3 0.3 0 0.3 0.3 0.3 0 Graz. Typical Plough 0.3 0.3 0 0 0.3 0.3 0 0 Graz. Rested Min. C 1.0 0.5 0.8 0.3 0.4 0.3 0.3 0.3

174

Appendix 2a-j: Mean values (± SE) for all response variables from Jealott’s Hill including reference existing grassland plots managed as low input EK2 grasslands.

Appendix 2a: Jealott’s Hill soil chemistry.

Soil carbon (g kg-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original Cut Typical n/a 24.07 26.40 28.77 0.89 1.73 0.99 Grassland Graz. Typical n/a 24.55 26.87 28.82 1.05 1.66 1.71

Grass Cut Typical Min. C 28.87 26.27 26.62 2.21 1.05 0.95 Cut Typical Plough 25.05 24.92 25.27 1.21 2.05 0.99 Graz. Typical Min. C 28.12 27.77 25.7 0.48 1.44 0.56 Graz. Typical Plough 24.70 25.50 26.8 1.10 0.60 1.62 GL Cut Typical Min. C 32.40 28.52 28.30 0.79 1.72 1.74 Cut Typical Plough 27.72 27.0 26.65 2.95 2.04 0.53 Graz. Typical Min. C 30.02 28.97 29.55 2.95 2.04 0.53 Graz. Typical Plough 26.02 25.8 29.17 3.62 2.03 1.26 GLF Cut Typical Min. C 29.92 27.65 28.70 2.87 1.50 2.85 Cut Typical Plough 28.85 26.35 26.52 3.07 1.84 1.81 Graz. Typical Min. C 26.45 28.07 27.90 1.07 1.30 2.18 Graz. Typical Plough 26.65 26.6 28.75 0.60 1.99 1.42

Soil nitrogen g kg-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original Cut Typical n/a 2.30 2.49 2.72 0.08 0.12 0.13 Grassland Graz. Typical n/a 2.25 2.54 2.72 0.11 0.10 0.09

Grass Cut Typical Min. C 2.64 2.50 2.47 0.19 0.05 0.08 Cut Typical Plough 2.37 2.35 2.35 0.09 0.12 0.09 Graz. Typical Min. C 2.60 2.57 2.42 0.03 0.09 0.05 Graz. Typical Plough 2.35 2.37 2.50 0.09 0.03 0.12 GL Cut Typical Min. C 3.00 2.65 2.62 0.09 0.12 0.19 Cut Typical Plough 2.61 2.57 2.45 0.21 0.17 0.14 Graz. Typical Min. C 2.85 2.71 2.80 0.21 0.17 0.14 Graz. Typical Plough 2.48 2.47 2.77 0.25 0.16 0.17 GLF Cut Typical Min. C 2.72 2.62 2.75 0.19 0.08 0.15 Cut Typical Plough 2.72 2.46 2.57 0.23 0.13 0.10 Graz. Typical Min. C 2.57 2.66 2.70 0.08 0.07 0.21 Graz. Typical Plough 2.59 2.54 2.75 0.07 0.12 0.13

175

Soil Olsen’s extractable phosphorus (mg kg-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original Cut Typical n/a 10.68 12.55 14.00 1.10 1.32 1.24 Grassland Graz. Typical n/a 11.35 13.20 15.00 0.90 0.85 4.42

Grass Cut Typical Min. C 14.84 13.10 12.50 1.21 1.25 1 Cut Typical Plough 13.72 15.55 15.00 1.04 2.83 1.05 Graz. Typical Min. C 17.04 17.40 13.75 2.74 2.11 2.07 Graz. Typical Plough 14.04 16.15 12.75 1.48 1.79 2.37 GL Cut Typical Min. C 23.03 18.35 11.00 1.05 1.58 1.19 Cut Typical Plough 14.95 14.75 11.75 0.91 2.54 2.42 Graz. Typical Min. C 15.31 15.25 12.75 0.91 2.54 2.42 Graz. Typical Plough 13.06 13.85 14.50 1.99 1.19 1.97 GL Cut Typical Min. C 14.80 11.90 12.75 1.26 1.37 2.95 Cut Typical Plough 14.48 11.80 11.50 0.76 0.78 1.73 Graz. Typical Min. C 15.12 11.55 11.50 2.10 1.03 0.74 Graz. Typical Plough 13.07 13.75 10.50 1.78 1.24 1.97

Total soil phosphorus (mg kg-1)

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original Cut Typical n/a 590.10 655.90 585.25 24.43 16.68 12.18 Grassland Graz. Typical n/a 626.52 721.22 624.5 19.71 53.33 33.14

Grass Cut Typical Min. C 738.47 686.62 597.0. 35.94 57.28 30.26 Cut Typical Plough 727.39 652.8 590.00 58.05 17.61 12.31 Graz. Typical Min. C 774.53 746.47 580.50 32.90 92.55 32.72 Graz. Typical Plough 696.12 638.07 577.50 11.61 38.12 29.31 GL Cut Typical Min. C 811.61 748.57 578.25 28.16 55.54 49.23 Cut Typical Plough 729.14 642.02 577.25 11.51 94.03 49.09 Graz. Typical Min. C 722.14 693.25 576.50 11.51 94.03 49.09 Graz. Typical Plough 766.70 676.27 617.25 58.21 56.99 53.77 GL Cut Typical Min. C 739.60 699.82 617.25 52.76 58.68 19.29 Cut Typical Plough 748.25 663.62 568.50 16.16 37.76 16.84 Graz. Typical Min. C 747.24 713.25 586.50 46.17 51.67 17.07 Graz. Typical Plough 680.55 670.92 576.25 27.87 11.33 29.33

176

Soil pH

Seed mix Man. Timing Cult. Means SE 2009 2010 2012 2009 2010 2012 Original Cut Typical n/a 6.40 6.50 6.97 0.12 0.07 0.19 Grassland Graz. Typical n/a 6.55 6.57 6.92 0.16 0.22 0.36

Grass Cut Typical Min. C 6.74 6.58 6.92 0.34 0.20 0.29 Cut Typical Plough 7.12 6.77 7.02 0.36 0.23 0.15 Graz. Typical Min. C 6.70 6.47 6.95 0.17 0.14 0.19 Graz. Typical Plough 6.92 6.69 7.10 0.33 0.16 0.23 GL Cut Typical Min. C 6.57 6.48 6.70 0.24 0.15 0.10 Cut Typical Plough 6.64 6.41 6.60 0.27 0.13 0.09 Graz. Typical Min. C 6.38 6.30 6.67 0.27 0.13 0.09 Graz. Typical Plough 6.75 6.59 6.60 0.37 0.19 0.16 GL Cut Typical Min. C 6.50 6.52 6.97 0.25 0.20 0.28 Cut Typical Plough 6.61 6.56 6.97 0.26 0.12 0.25 Graz. Typical Min. C 6.68 6.44 6.75 0.38 0.18 0.15 Graz. Typical Plough 6.82 6.58 6.72 0.23 0.04 0.02

PLFA Fungal to bacterial ratio

Seed mix Man. Timing Cult. Means SE 2012 2012 Original grassland Cut Typical n/a 0.0903 0.0072 Graz. Typical n/a 0.0652 0.0110 Grass Cut Rest. Plough 0.0782 0.0161 Cut Rest. Min. Cult. 0.0686 0.0096 Graz. Typical. Plough 0.0839 0.0113 Graz. Typical. Min. Cult. 0.0768 0.0157 GL Cut Rest. Plough 0.0635 0.0030 Cut Rest. Min. Cult. 0.1064 0.0252 Graz. Typical. Plough 0.0586 0.0026 Graz. Typical. Min. Cult. 0.0751 0.0163 GLF Cut Rest. Plough 0.0808 0.0218 Cut Rest. Min. Cult. 0.1225 0.0623 Graz. Typical. Plough 0.0599 0.0029 Graz. Typical. Min. Cult. 0.0600 0.0034

177

Appendix 2b: Jealott’s Hill herbage production from combined yearly silage cuts (tonnes ha-1).

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 3.88 2.89 8.19 2.97 0.65 0.73 0.73 0.26 Grass Cut Rest. Plough 1.86 0.43 1.81 0.66 0.34 0.06 0.16 0.09 Cut Rest. Min. Cult. 2.60 0.49 2.09 0.71 0.72 0.13 0.37 0.09 Cut Typical. Plough 1.92 0.84 2.33 0.75 0.32 0.09 0.36 0.11 Cut Typical. Min. Cult. 5.32 1.01 2.56 0.90 1.78 0.20 0.12 0.12 GL Cut Rest. Plough 7.33 2.41 5.29 2.84 0.98 0.81 1.92 0.19 Cut Rest. Min. Cult. 8.34 2.50 8.01 3.34 1.34 0.58 0.83 0.39 Cut Typical. Plough 8.23 3.86 5.47 2.66 0.52 0.85 2.01 0.22 Cut Typical. Min. Cult. 8.07 3.65 4.46 3.24 0.75 1.02 2.23 0.62 GLF Cut Rest. Plough 8.67 2.69 8.84 3.11 0.32 0.34 0.49 0.44 Cut Rest. Min. Cult. 7.86 2.06 9.26 3.17 0.37 0.33 1.58 0.51 Cut Typical. Plough 8.40 4.54 9.21 2.68 0.17 0.37 2.47 0.33 Cut Typical. Min. Cult. 8.45 6.32 10.59 2.50 0.57 2.88 0.64 0.24

178

Appendix 2c: Jealott’s Hill herbage forage quality.

Herbage Nitrogen (% w/w)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 2.05 2.27 1.62 1.24 0.13 0.05 0.19 0.17 Grass Cut Rest. Plough 1.15 1.56 1.11 1.04 0.01 0.05 0.05 0.35 Cut Rest. Min. Cult. 1.28 1.50 1.20 1.50 0.02 0.09 0.07 0.19 Cut Typical. Plough 1.12 1.50 1.11 1.30 0.04 0.03 0.06 0.10 Cut Typical. Min. Cult. 1.40 1.58 1.11 1.22 0.05 0.03 0.01 0.10 GL Cut Rest. Plough 2.12 2.01 1.03 0.90 0.06 0.13 0.35 0.11 Cut Rest. Min. Cult. 2.30 2.13 1.52 0.72 0.07 0.09 0.10 0.06 Cut Typical. Plough 1.94 1.83 1.57 0.83 0.16 0.12 0.09 0.03 Cut Typical. Min. Cult. 2.20 2.40 1.05 0.77 0.09 0.32 0.37 0.05 GLF Cut Rest. Plough 2.26 2.02 1.30 0.93 0.10 0.06 0.15 0.01 Cut Rest. Min. Cult. 2.17 1.82 1.36 0.88 0.05 0.12 0.06 0.04 Cut Typical. Plough 2.31 2.02 1.47 0.85 0.33 0.21 0.06 0.03 Cut Typical. Min. Cult. 2.32 2.35 1.46 0.93 0.08 0.13 0.06 0.04

Herbage calcium (% w/w)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.83 0.89 0.71 0.53 0.07 0.13 0.05 0.06 Grass Cut Rest. Plough 0.46 0.53 0.50 0.41 0.01 0.04 0.06 0.14 Cut Rest. Min. Cult. 0.57 0.52 0.50 0.77 0.05 0.02 0.06 0.12 Cut Typical. Plough 0.45 0.53 0.59 0.51 0.02 0.05 0.05 0.05 Cut Typical. Min. Cult. 0.61 0.49 0.43 0.53 0.04 0.02 0.02 0.05 GL Cut Rest. Plough 0.85 0.69 0.34 0.34 0.07 0.06 0.12 0.06 Cut Rest. Min. Cult. 0.90 0.80 0.52 0.30 0.03 0.06 0.03 0.04 Cut Typical. Plough 0.80 0.86 0.63 0.36 0.09 0.10 0.02 0.06 Cut Typical. Min. Cult. 0.72 0.88 0.41 0.32 0.08 0.07 0.16 0.01 GLF Cut Rest. Plough 0.98 0.93 0.71 0.63 0.07 0.09 0.03 0.08 Cut Rest. Min. Cult. 0.87 0.75 0.55 0.64 0.04 0.13 0.07 0.09 Cut Typical. Plough 0.96 0.98 0.91 0.58 0.12 0.08 0.07 0.03 Cut Typical. Min. Cult. 0.93 1.07 0.62 0.50 0.03 0.05 0.09 0.04

179

Herbage potassium (% w/w)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 2.16 2.39 1.72 1.33 0.14 0.18 0.11 0.14 Grass Cut Rest. Plough 1.66 2.03 1.66 1.16 0.03 0.09 0.07 0.41 Cut Rest. Min. Cult. 1.69 2.09 1.52 1.47 0.04 0.09 0.07 0.13 Cut Typical. Plough 1.69 1.86 1.73 1.36 0.10 0.06 0.09 0.07 Cut Typical. Min. Cult. 1.84 1.94 1.56 1.44 0.05 0.03 0.06 0.17 GL Cut Rest. Plough 2.26 2.16 1.11 1.06 0.18 0.13 0.37 0.08 Cut Rest. Min. Cult. 2.43 2.24 1.56 0.85 0.10 0.07 0.06 0.06 Cut Typical. Plough 2.18 2.21 1.72 1.06 0.22 0.05 0.06 0.03 Cut Typical. Min. Cult. 2.23 2.48 1.19 0.93 0.16 0.18 0.40 0.06 GLF Cut Rest. Plough 2.42 2.45 2.00 1.52 0.10 0.10 0.10 0.04 Cut Rest. Min. Cult. 2.30 2.19 1.57 1.43 0.06 0.20 0.07 0.07 Cut Typical. Plough 2.60 2.34 2.02 1.43 0.22 0.12 0.06 0.09 Cut Typical. Min. Cult. 2.28 2.50 1.84 1.29 0.12 0.15 0.05 0.14

Herbage magnesium (% w/w) Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.17 0.19 0.17 0.13 0.02 0.03 0.02 0.01 Grass Cut Rest. Plough 0.11 0.14 0.14 0.10 0.01 0.01 0.02 0.04 Cut Rest. Min. Cult. 0.12 0.13 0.13 0.16 0.01 0.01 0.01 0.01 Cut Typical. Plough 0.12 0.14 0.16 0.13 0.01 0.01 0.02 0.01 Cut Typical. Min. Cult. 0.13 0.13 0.12 0.13 0.01 0.01 0.01 0.02 GL Cut Rest. Plough 0.22 0.19 0.13 0.11 0.01 0.01 0.04 0.01 Cut Rest. Min. Cult. 0.22 0.21 0.18 0.09 0.01 0.02 0.02 0.01 Cut Typical. Plough 0.18 0.21 0.17 0.11 0.01 0.02 0.01 0.01 Cut Typical. Min. Cult. 0.18 0.21 0.13 0.10 0.01 0.02 0.06 0.00 GLF Cut Rest. Plough 0.21 0.21 0.18 0.16 0.00 0.01 0.01 0.01 Cut Rest. Min. Cult. 0.19 0.17 0.15 0.17 0.01 0.02 0.02 0.01 Cut Typical. Plough 0.23 0.24 0.24 0.14 0.02 0.02 0.01 0.01 Cut Typical. Min. Cult. 0.21 0.23 0.18 0.12 0.01 0.02 0.01 0.01

180

Herbage sodium (% w/w) Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.09 0.15 0.22 0.05 0.01 0.02 0.03 0.01 Grass Cut Rest. Plough 0.03 0.04 0.04 0.03 0.00 0.01 0.00 0.01 Cut Rest. Min. Cult. 0.06 0.05 0.06 0.05 0.00 0.01 0.01 0.00 Cut Typical. Plough 0.04 0.07 0.08 0.04 0.01 0.03 0.03 0.01 Cut Typical. Min. Cult. 0.07 0.06 0.05 0.04 0.01 0.01 0.00 0.00 GL Cut Rest. Plough 0.06 0.06 0.07 0.03 0.01 0.01 0.02 0.00 Cut Rest. Min. Cult. 0.09 0.10 0.13 0.03 0.01 0.01 0.02 0.00 Cut Typical. Plough 0.11 0.09 0.09 0.02 0.03 0.02 0.02 0.00 Cut Typical. Min. Cult. 0.09 0.11 0.08 0.02 0.01 0.01 0.05 0.00 GLF Cut Rest. Plough 0.07 0.09 0.08 0.05 0.01 0.02 0.01 0.01 Cut Rest. Min. Cult. 0.10 0.08 0.11 0.05 0.02 0.02 0.03 0.01 Cut Typical. Plough 0.07 0.10 0.17 0.05 0.01 0.01 0.02 0.01 Cut Typical. Min. Cult. 0.09 0.10 0.12 0.04 0.02 0.02 0.01 0.00

Herbage phosphorus (% w/w)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.20 0.25 0.18 0.18 0.01 0.00 0.00 0.02 Grass Cut Rest. Plough 0.18 0.26 0.22 0.16 0.01 0.02 0.01 0.05 Cut Rest. Min. Cult. 0.21 0.24 0.19 0.19 0.01 0.01 0.01 0.01 Cut Typical. Plough 0.17 0.25 0.21 0.19 0.01 0.00 0.01 0.01 Cut Typical. Min. Cult. 0.20 0.23 0.21 0.20 0.01 0.01 0.01 0.02 GL Cut Rest. Plough 0.21 0.23 0.13 0.13 0.01 0.01 0.04 0.02 Cut Rest. Min. Cult. 0.22 0.24 0.18 0.11 0.01 0.01 0.00 0.01 Cut Typical. Plough 0.20 0.23 0.17 0.12 0.02 0.01 0.00 0.00 Cut Typical. Min. Cult. 0.22 0.24 0.13 0.12 0.01 0.02 0.04 0.00 GLF Cut Rest. Plough 0.20 0.22 0.18 0.18 0.01 0.00 0.01 0.01 Cut Rest. Min. Cult. 0.23 0.24 0.20 0.17 0.01 0.01 0.01 0.01 Cut Typical. Plough 0.25 0.26 0.22 0.17 0.04 0.02 0.01 0.01 Cut Typical. Min. Cult. 0.22 0.26 0.19 0.15 0.01 0.01 0.00 0.01

181

DOMD (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 66.00 65.61 65.83 51.11 0.08 1.07 0.70 1.93 Grass Cut Rest. Plough 65.87 62.93 61.90 50.54 0.29 0.69 1.37 2.18 Cut Rest. Min. Cult. 65.38 63.16 61.50 52.38 0.64 0.61 1.58 1.35 Cut Typical. Plough 64.83 62.43 62.91 51.63 0.62 1.46 0.68 0.69 Cut Typical. Min. Cult. 65.57 65.19 63.63 50.49 1.04 1.19 1.06 1.29 GL Cut Rest. Plough 65.00 61.53 60.50 46.61 0.78 0.61 1.59 0.79 Cut Rest. Min. Cult. 66.82 64.82 63.19 44.76 0.33 1.74 1.68 0.14 Cut Typical. Plough 65.01 65.36 62.44 45.71 0.51 1.16 1.83 0.96 Cut Typical. Min. Cult. 65.55 65.36 64.01 45.25 1.35 0.83 1.23 1.07 GLF Cut Rest. Plough 65.66 63.14 57.27 47.12 0.76 0.52 1.57 1.00 Cut Rest. Min. Cult. 67.16 62.77 60.66 46.70 0.66 1.11 1.97 0.97 Cut Typical. Plough 63.69 64.24 62.06 45.68 0.74 1.39 0.48 0.33 Cut Typical. Min. Cult. 68.00 64.72 61.72 46.28 0.28 0.90 1.21 0.66

182

Appendix 2d: Jealott’s Hill summed percentage cover of grasses, legumes and non-legume forbs.

Grass summed percentage cover (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 34.1 47.9 41.4 86.2 7.4 3.4 4.6 5.4 Graz. Typical n/a 30.0 45.6 42.9 40.7 2.3 6.2 8.5 5.1 Grass Cut Rest. Plough 60.5 45.1 61.7 84.2 2.5 2.2 10.1 3.4 Cut Rest. Min. Cult. 75.6 51.6 72.3 86.1 4.7 2.0 5.0 3.0 Cut Typical. Plough 61.3 48.5 70.6 84.4 3.6 6.2 3.1 3.8 Cut Typical. Min. Cult. 79.8 55.2 68.1 81.0 5.2 7.4 5.2 5.0 Graz. Rest. Plough 69.3 62.5 73.0 79.4 6.4 8.7 10.1 9.9 Graz. Rest. Min. Cult. 72.7 63.4 78.9 71.6 4.2 7.7 3.2 4.5 Graz. Typical. Plough 56.6 49.2 62.6 69.7 2.3 4.4 5.0 3.1 Graz. Typical. Min. Cult. 73.6 55.1 65.7 63.9 4.5 4.3 3.3 6.1 GL Cut Rest. Plough 33.1 50.9 36.8 65.6 5.6 2.1 3.0 16.8 Cut Rest. Min. Cult. 38.1 53.3 48.3 80.5 7.8 6.6 8.2 1.7 Cut Typical. Plough 22.6 31.3 40.9 81.8 9.7 5.6 6.2 6.8 Cut Typical. Min. Cult. 32.1 47.2 45.9 60.4 5.1 4.4 5.3 12.7 Graz. Rest. Plough 37.9 49.2 55.6 40.0 6.1 4.7 7.9 15.6 Graz. Rest. Min. Cult. 40.9 53.3 52.9 38.9 5.1 4.8 6.4 14.1 Graz. Typical. Plough 33.4 26.1 46.3 56.7 0.7 2.8 10.1 13.2 Graz. Typical. Min. Cult. 35.9 41.5 52.6 52.5 1.6 5.5 4.3 12.4 GLF Cut Rest. Plough 19.3 20.6 34.3 53.0 7.7 2.6 4.4 2.6 Cut Rest. Min. Cult. 37.6 45.8 22.4 60.0 12.5 3.3 7.9 6.7 Cut Typical. Plough 25.4 15.8 30.2 59.8 9.7 1.5 3.0 8.0 Cut Typical. Min. Cult. 32.5 33.1 29.4 48.0 7.1 6.3 4.3 3.3 Graz. Rest. Plough 22.5 29.2 56.8 45.0 2.6 2.9 7.3 6.0 Graz. Rest. Min. Cult. 32.7 37.7 54.9 39.0 1.7 6.1 4.5 3.4 Graz. Typical. Plough 25.8 25.4 57.5 58.6 2.9 3.7 3.6 8.2 Graz. Typical. Min. Cult. 25.7 30.4 48.9 45.8 6.5 3.7 8.4 5.2

Non-legume forb summed percentage cover (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.0 0.0 0.3 0.7 0.0 0.0 0.3 0.5 Graz. Typical n/a 0.0 0.0 0.1 0.2 0.0 0.0 0.1 0.1 Grass Cut Rest. Plough 0.3 0.2 2.6 1.2 0.3 0.2 2.0 1.0 Cut Rest. Min. Cult. 0.0 0.1 0.0 0.8 0.0 0.1 0.0 0.3 Cut Typical. Plough 0.3 0.0 0.0 1.6 0.2 0.0 0.0 0.8 Cut Typical. Min. Cult. 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.2 Graz. Rest. Plough 0.2 0.2 3.9 7.0 0.1 0.2 3.5 5.8 Graz. Rest. Min. Cult. 0.0 0.1 0.4 1.2 0.0 0.1 0.2 0.7 Graz. Typical. Plough 0.1 0.1 0.0 2.4 0.1 0.1 0.0 0.6 Graz. Typical. Min. Cult. 0.0 0.1 0.0 1.7 0.0 0.1 0.0 1.1 GL Cut Rest. Plough 0.2 0.2 0.0 6.7 0.2 0.1 0.0 2.6 Cut Rest. Min. Cult. 0.1 0.5 0.0 2.6 0.1 0.4 0.0 0.9 Cut Typical. Plough 0.5 0.0 0.3 3.6 0.2 0.0 0.2 3.3 Cut Typical. Min. Cult. 0.4 0.2 0.3 1.5 0.4 0.2 0.2 1.5 Graz. Rest. Plough 0.5 0.5 5.0 2.7 0.5 0.4 4.1 1.1 Graz. Rest. Min. Cult. 0.2 0.3 0.1 1.4 0.2 0.2 0.1 0.8 Graz. Typical. Plough 0.2 0.4 0.1 2.5 0.1 0.1 0.1 1.4 Graz. Typical. Min. Cult. 0.0 0.1 0.1 0.5 0.0 0.0 0.1 0.3 GLF Cut Rest. Plough 22.8 23.1 26.5 50.6 4.2 1.4 0.6 3.9 Cut Rest. Min. Cult. 13.3 12.3 16.9 49.7 3.2 3.2 6.3 5.8 Cut Typical. Plough 25.9 19.9 25.1 40.9 5.5 3.5 5.6 6.5 Cut Typical. Min. Cult. 17.6 16.6 20.2 53.4 2.8 5.4 2.6 7.9 Graz. Rest. Plough 23.3 30.9 25.5 39.7 6.5 8.2 3.4 5.0 Graz. Rest. Min. Cult. 10.1 20.4 21.7 20.6 4.2 4.8 4.1 5.8 Graz. Typical. Plough 19.6 12.3 15.9 26.9 5.8 2.7 0.7 3.7 Graz. Typical. Min. Cult. 5.7 5.4 6.1 20.5 1.2 1.6 1.1 3.4

183

Legume summed percentage cover (%)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 75.2 38.3 54.8 18.9 7.9 4.7 4.2 5.3 Graz. Typical n/a 60.3 17.0 31.3 66.8 7.7 4.9 4.6 6.5 Grass Cut Rest. Plough 0.0 0.3 16.2 12.4 0.0 0.3 10.1 5.7 Cut Rest. Min. Cult. 0.0 0.4 3.4 11.9 0.0 0.3 2.2 3.6 Cut Typical. Plough 0.7 0.3 4.0 12.9 0.5 0.2 1.6 4.3 Cut Typical. Min. Cult. 0.0 0.1 9.4 27.9 0.0 0.1 6.1 8.6 Graz. Rest. Plough 0.0 0.1 2.7 13.4 0.0 0.1 0.9 5.9 Graz. Rest. Min. Cult. 2.5 0.3 6.1 22.1 1.4 0.2 2.3 9.1 Graz. Typical. Plough 0.0 0.1 2.2 34.5 0.0 0.1 1.1 6.6 Graz. Typical. Min. Cult. 0.8 0.3 4.6 36.6 0.8 0.1 1.9 8.1 GL Cut Rest. Plough 80.7 27.2 59.6 34.0 4.5 4.0 2.7 17.6 Cut Rest. Min. Cult. 83.7 32.5 48.9 29.9 5.2 4.9 9.8 8.0 Cut Typical. Plough 87.8 54.0 57.1 23.5 9.9 9.5 7.0 9.2 Cut Typical. Min. Cult. 80.7 41.9 51.4 44.4 3.8 5.6 4.3 12.4 Graz. Rest. Plough 61.2 53.6 33.0 62.4 10.3 15.3 4.3 17.5 Graz. Rest. Min. Cult. 61.9 39.6 42.6 67.1 10.3 5.5 7.0 15.2 Graz. Typical. Plough 49.9 30.1 27.5 45.4 9.2 3.8 9.6 15.3 Graz. Typical. Min. Cult. 45.8 22.1 29.1 49.6 11.2 2.3 5.5 13.1 GLF Cut Rest. Plough 85.0 20.6 31.7 2.7 2.4 4.6 3.1 1.0 Cut Rest. Min. Cult. 80.0 20.3 30.2 3.9 4.3 2.1 11.1 3.1 Cut Typical. Plough 78.3 45.7 44.2 5.1 3.9 1.0 6.3 2.0 Cut Typical. Min. Cult. 72.8 35.1 44.2 9.0 1.8 2.3 4.7 2.0 Graz. Rest. Plough 60.3 29.8 12.0 17.7 3.9 2.8 8.1 4.1 Graz. Rest. Min. Cult. 67.6 32.2 20.4 46.6 4.5 2.8 5.6 7.8 Graz. Typical. Plough 48.6 24.5 7.9 21.9 8.8 4.2 2.4 13.2 Graz. Typical. Min. Cult. 65.3 18.6 24.9 36.6 14.5 3.8 5.9 15.1

184

Appendix 2e: Jealott’s Hill species richness of grasses, legumes and non-legume forbs.

Grass species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 1.0 1.0 1.0 1.0 0.0 0.0 0.0 0.0 Graz. Typical n/a 1.0 1.0 1.0 1.0 0.0 0.0 0.0 0.0 Grass Cut Rest. Plough 0.0 0.3 1.5 2.0 0.0 0.3 0.6 0.7 Cut Rest. Min. Cult. 0.0 0.5 1.3 1.8 0.0 0.3 0.5 0.3 Cut Typical. Plough 0.8 0.5 1.3 1.3 0.3 0.3 0.3 0.3 Cut Typical. Min. Cult. 0.0 0.5 2.0 1.3 0.0 0.3 0.7 0.3 Graz. Rest. Plough 0.0 0.3 1.5 1.0 0.0 0.3 0.5 0.4 Graz. Rest. Min. Cult. 0.8 0.5 1.0 1.3 0.5 0.3 0.4 0.6 Graz. Typical. Plough 0.0 0.3 1.0 1.3 0.0 0.3 0.0 0.3 Graz. Typical. Min. Cult. 0.3 0.8 1.0 1.8 0.3 0.3 0.0 0.5 GL Cut Rest. Plough 5.5 4.5 4.8 3.5 0.6 0.6 0.5 0.3 Cut Rest. Min. Cult. 4.5 3.3 3.3 3.3 0.5 0.3 0.3 0.3 Cut Typical. Plough 4.3 4.0 3.5 2.8 0.5 0.4 0.3 0.3 Cut Typical. Min. Cult. 4.8 3.8 3.3 3.5 0.8 0.3 0.3 0.3 Graz. Rest. Plough 4.8 4.8 3.8 3.0 0.3 0.3 0.3 0.7 Graz. Rest. Min. Cult. 4.0 3.8 3.5 3.0 0.0 0.3 0.3 0.4 Graz. Typical. Plough 5.0 4.0 3.0 2.5 0.7 0.0 0.4 0.3 Graz. Typical. Min. Cult. 3.5 3.8 2.5 2.0 0.3 0.3 0.3 0.4 GLF Cut Rest. Plough 4.0 4.3 4.0 1.0 0.4 0.3 0.4 0.0 Cut Rest. Min. Cult. 4.3 4.0 2.3 1.5 0.3 0.0 0.8 0.6 Cut Typical. Plough 4.3 3.5 4.5 2.3 0.6 0.3 0.3 0.3 Cut Typical. Min. Cult. 3.8 3.3 3.3 3.0 0.3 0.3 0.3 0.4 Graz. Rest. Plough 4.5 4.8 2.3 2.8 0.3 0.3 1.1 0.5 Graz. Rest. Min. Cult. 4.0 3.5 3.3 3.0 0.0 0.3 0.3 0.4 Graz. Typical. Plough 4.5 4.0 2.3 1.8 0.3 0.6 0.5 0.8 Graz. Typical. Min. Cult. 3.8 3.5 2.0 1.8 0.6 0.3 0.4 0.6

Non-legume forb species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 1.0 0.8 1.3 1.5 0.0 0.3 0.3 0.3 Graz. Typical n/a 0.8 0.8 0.8 1.5 0.3 0.3 0.5 0.3 Grass Cut Rest. Plough 1.5 1.3 2.0 2.0 0.5 0.3 0.7 1.1 Cut Rest. Min. Cult. 1.0 1.3 0.5 2.3 0.0 0.3 0.3 0.3 Cut Typical. Plough 2.0 1.3 0.8 2.5 0.7 0.3 0.3 0.5 Cut Typical. Min. Cult. 1.3 1.3 0.5 1.5 0.3 0.3 0.3 0.3 Graz. Rest. Plough 1.8 1.5 2.3 2.8 0.3 0.5 0.6 1.0 Graz. Rest. Min. Cult. 1.0 2.0 2.0 2.3 0.0 1.0 0.4 1.0 Graz. Typical. Plough 1.5 2.0 1.0 2.8 0.5 0.4 0.0 0.6 Graz. Typical. Min. Cult. 0.8 1.3 0.8 2.5 0.3 0.3 0.3 0.6 GL Cut Rest. Plough 1.5 1.5 1.0 2.5 0.5 0.3 0.0 1.0 Cut Rest. Min. Cult. 1.3 1.8 1.0 2.5 0.3 0.5 0.0 0.3 Cut Typical. Plough 2.0 1.0 1.5 2.5 0.0 0.0 0.6 1.0 Cut Typical. Min. Cult. 1.3 1.3 1.5 1.3 0.3 0.3 0.3 0.3 Graz. Rest. Plough 1.5 1.5 3.3 2.3 0.9 0.6 0.9 0.5 Graz. Rest. Min. Cult. 1.0 1.5 1.3 2.5 0.4 0.3 0.3 0.6 Graz. Typical. Plough 2.3 2.5 1.0 2.3 0.6 0.3 0.4 0.6 Graz. Typical. Min. Cult. 0.8 1.3 1.0 2.0 0.3 0.5 0.4 0.4 GLF Cut Rest. Plough 4.5 5.5 5.0 6.0 0.3 0.3 0.4 0.7 Cut Rest. Min. Cult. 5.5 4.0 2.8 5.3 0.6 0.4 0.9 0.8 Cut Typical. Plough 4.8 5.0 4.5 6.8 0.6 0.4 0.6 0.9 Cut Typical. Min. Cult. 5.0 4.8 4.8 5.0 0.4 0.8 0.5 0.8 Graz. Rest. Plough 5.5 6.0 5.5 6.3 0.6 0.4 0.6 0.3 Graz. Rest. Min. Cult. 4.8 5.5 4.5 3.0 0.3 0.3 0.9 0.7 Graz. Typical. Plough 5.0 6.8 4.8 4.3 0.4 0.9 0.3 0.8 Graz. Typical. Min. Cult. 5.0 5.8 4.3 3.8 0.0 0.3 0.3 0.6

185

Legume species richness

186

Appendix 2f: Jealott’s Hill density of legumes and non-legume forb flower heads.

Non-legume forb flower head density (m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Graz. Typical n/a 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Grass Cut Rest. Plough 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Cut Rest. Min. Cult. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Cut Typical. Plough 0.0 0.0 0.4 0.0 0.0 0.0 0.4 0.0 Cut Typical. Min. Cult. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Graz. Rest. Plough 0.0 0.2 0.5 0.0 0.0 0.2 0.4 0.0 Graz. Rest. Min. Cult. 0.0 0.0 0.8 0.1 0.0 0.0 0.4 0.1 Graz. Typical. Plough 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Graz. Typical. Min. Cult. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 GL Cut Rest. Plough 0.0 0.3 13.4 0.4 0.0 0.3 2.3 0.2 Cut Rest. Min. Cult. 0.0 0.2 5.8 0.0 0.0 0.2 3.0 0.0 Cut Typical. Plough 0.3 0.1 7.1 0.1 0.3 0.1 2.8 0.1 Cut Typical. Min. Cult. 0.0 0.0 7.0 0.0 0.0 0.0 2.0 0.0 Graz. Rest. Plough 0.2 0.0 14.5 0.5 0.2 0.0 7.2 0.2 Graz. Rest. Min. Cult. 0.2 0.2 7.9 0.1 0.2 0.2 4.6 0.1 Graz. Typical. Plough 0.0 0.0 0.2 0.0 0.0 0.0 0.1 0.0 Graz. Typical. Min. Cult. 0.0 0.0 0.3 0.2 0.0 0.0 0.2 0.2 GLF Cut Rest. Plough 7.1 4.0 14.0 19.7 1.9 0.8 3.7 1.0 Cut Rest. Min. Cult. 1.8 1.9 10.5 22.0 0.5 0.8 2.1 1.4 Cut Typical. Plough 3.0 2.7 13.5 20.8 1.6 0.7 2.6 1.0 Cut Typical. Min. Cult. 0.5 2.5 6.5 21.7 0.2 0.6 1.4 0.7 Graz. Rest. Plough 1.8 2.5 10.3 10.8 0.7 0.6 1.4 3.1 Graz. Rest. Min. Cult. 1.4 1.1 8.8 8.7 0.5 0.5 0.8 2.3 Graz. Typical. Plough 0.6 0.8 5.4 5.8 0.2 0.2 1.3 1.5 Graz. Typical. Min. Cult. 0.4 0.5 2.0 4.2 0.2 0.5 0.4 0.3

Legume flower head density (m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 15.4 46.5 45.9 1.8 5.0 2.4 1.5 0.8 Graz. Typical n/a 7.8 21.5 35.9 14.8 2.9 3.1 5.7 5.1 Grass Cut Rest. Plough 0.0 0.0 3.0 1.7 0.0 0.0 1.6 0.6 Cut Rest. Min. Cult. 0.0 0.0 2.2 4.1 0.0 0.0 1.7 0.6 Cut Typical. Plough 0.0 0.1 3.3 1.0 0.0 0.1 0.5 0.6 Cut Typical. Min. Cult. 0.0 0.0 6.9 4.3 0.0 0.0 4.0 1.7 Graz. Rest. Plough 1.7 0.0 2.1 2.1 1.6 0.0 0.8 0.5 Graz. Rest. Min. Cult. 0.1 1.3 8.7 4.2 0.1 1.3 5.4 2.6 Graz. Typical. Plough 0.0 0.0 3.3 5.2 0.0 0.0 0.9 1.1 Graz. Typical. Min. Cult. 0.0 0.0 3.6 7.5 0.0 0.0 2.0 3.6 GL Cut Rest. Plough 85.7 26.7 20.4 4.1 5.6 2.3 3.4 1.3 Cut Rest. Min. Cult. 81.0 34.2 31.8 5.8 10.4 4.4 5.5 1.7 Cut Typical. Plough 70.3 55.7 36.6 3.8 9.0 12.2 2.9 1.2 Cut Typical. Min. Cult. 59.3 52.3 43.1 3.5 3.1 4.6 6.5 1.1 Graz. Rest. Plough 34.0 32.8 16.0 13.1 15.7 5.6 3.4 5.0 Graz. Rest. Min. Cult. 30.5 35.3 21.9 11.2 10.5 2.5 8.3 3.9 Graz. Typical. Plough 12.2 21.5 24.2 15.4 2.5 3.8 5.0 3.8 Graz. Typical. Min. Cult. 13.2 14.3 21.0 11.5 3.5 2.7 5.6 2.7 GLF Cut Rest. Plough 83.5 21.1 9.6 1.7 9.1 8.1 0.6 0.8 Cut Rest. Min. Cult. 58.3 37.8 27.1 2.0 8.7 10.0 6.7 0.7 Cut Typical. Plough 58.3 35.9 34.1 2.3 9.1 3.5 3.6 0.6 Cut Typical. Min. Cult. 54.5 40.6 39.2 2.3 7.3 2.4 4.5 0.6 Graz. Rest. Plough 27.0 24.5 10.8 8.0 8.9 8.7 4.7 1.9 Graz. Rest. Min. Cult. 28.3 17.3 18.0 10.5 8.5 4.1 4.8 3.2 Graz. Typical. Plough 12.0 11.9 9.5 6.2 3.5 2.5 2.4 1.7 Graz. Typical. Min. Cult. 15.4 17.0 15.6 8.7 3.9 1.8 2.3 1.8

187

Appendix 2g: Jealott’s Hill density of grasses, legumes and non-legume forb winter seed heads.

Winter grass seed head density (seed heads m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.5 0.0 0.7 0.3 0.5 0.0 0.7 0.3 Graz. Typical n/a 2.0 0.0 4.5 3.5 1.2 0.0 1.1 2.1 Grass Cut Rest. Plough 0.0 0.0 2.7 1.5 0.0 0.0 0.8 0.9 Cut Rest. Min. Cult. 0.0 0.0 0.3 2.2 0.0 0.0 0.3 1.5 Cut Typical. Plough 0.0 0.0 1.2 0.2 0.0 0.0 1.0 0.2 Cut Typical. Min. Cult. 0.0 0.0 0.0 0.8 0.0 0.0 0.0 0.8 Graz. Rest. Plough 0.0 0.0 0.3 1.7 0.0 0.0 0.3 1.0 Graz. Rest. Min. Cult. 0.7 0.0 0.2 0.5 0.7 0.0 0.2 0.5 Graz. Typical. Plough 0.0 0.0 1.2 0.0 0.0 0.0 0.4 0.0 Graz. Typical. Min. Cult. 0.2 0.0 2.3 0.8 0.2 0.0 1.0 0.8 GL Cut Rest. Plough 0.0 81.0 47.5 1.3 0.0 32.7 16.6 0.7 Cut Rest. Min. Cult. 0.2 48.3 29.8 2.2 0.2 17.1 11.7 1.1 Cut Typical. Plough 0.2 0.0 1.5 0.5 0.2 0.0 0.9 0.5 Cut Typical. Min. Cult. 0.0 0.0 2.2 2.5 0.0 0.0 0.8 0.7 Graz. Rest. Plough 81.3 77.7 4.3 1.5 58.6 27.5 1.5 1.3 Graz. Rest. Min. Cult. 47.7 33.8 2.8 0.8 41.6 14.3 0.7 0.6 Graz. Typical. Plough 0.3 1.0 7.8 3.7 0.3 1.0 2.6 1.6 Graz. Typical. Min. Cult. 2.2 0.0 4.3 1.3 0.6 0.0 1.7 0.8 GLF Cut Rest. Plough 1.2 44.8 34.3 0.0 0.6 21.9 5.1 0.0 Cut Rest. Min. Cult. 3.0 32.7 43.8 0.2 3.0 17.0 9.6 0.2 Cut Typical. Plough 0.2 6.7 3.2 1.8 0.2 6.7 1.1 1.4 Cut Typical. Min. Cult. 0.0 21.7 3.3 0.5 0.0 21.4 0.4 0.3 Graz. Rest. Plough 16.8 35.8 6.3 1.7 4.6 23.1 2.7 1.0 Graz. Rest. Min. Cult. 11.7 71.8 5.0 1.7 6.3 39.2 2.1 1.0 Graz. Typical. Plough 2.3 0.5 6.8 0.5 1.0 0.5 2.5 0.5 Graz. Typical. Min. Cult. 5.5 0.0 4.7 0.5 1.8 0.0 1.2 0.3

Winter non-legume forb seed head density (seed heads m-2)

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.0 0.0 0.0 1.0 0.0 0.0 0.0 1.0 Graz. Typical n/a 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.2 Grass Cut Rest. Plough 0.0 0.0 0.0 0.8 0.0 0.0 0.0 0.8 Cut Rest. Min. Cult. 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.3 Cut Typical. Plough 0.0 0.0 0.0 1.5 0.0 0.0 0.0 1.5 Cut Typical. Min. Cult. 0.0 0.0 0.0 0.5 0.0 0.0 0.0 0.5 Graz. Rest. Plough 0.0 0.0 0.0 1.0 0.0 0.0 0.0 1.0 Graz. Rest. Min. Cult. 3.5 0.0 0.0 2.0 3.5 0.0 0.0 2.0 Graz. Typical. Plough 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.2 Graz. Typical. Min. Cult. 1.0 0.0 0.0 1.2 1.0 0.0 0.0 1.2 GL Cut Rest. Plough 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.3 Cut Rest. Min. Cult. 0.0 0.0 0.0 1.5 0.0 0.0 0.0 1.3 Cut Typical. Plough 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Cut Typical. Min. Cult. 0.0 0.0 0.0 0.8 0.0 0.0 0.0 0.8 Graz. Rest. Plough 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Graz. Rest. Min. Cult. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Graz. Typical. Plough 0.0 0.0 0.0 1.3 0.0 0.0 0.0 1.3 Graz. Typical. Min. Cult. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 GLF Cut Rest. Plough 0.0 217.8 38.5 0.0 0.0 87.6 10.5 0.0 Cut Rest. Min. Cult. 1.8 175.3 47.5 0.0 1.8 94.7 8.5 0.0 Cut Typical. Plough 0.0 44.0 0.0 0.0 0.0 44.0 0.0 0.0 Cut Typical. Min. Cult. 1.0 63.7 0.0 0.0 1.0 52.0 0.0 0.0 Graz. Rest. Plough 100.0 457.8 15.0 0.0 34.0 99.7 6.0 0.0 Graz. Rest. Min. Cult. 37.5 368.3 9.3 0.2 19.3 64.6 2.6 0.2 Graz. Typical. Plough 17.5 108.0 11.2 0.0 4.8 17.8 5.9 0.0 Graz. Typical. Min. Cult. 10.8 45.0 6.5 0.0 4.0 5.5 4.4 0.0

188

Winter legume seed head density (seed heads m-2)

189

Appendix 2h: Jealott’s Hill beetle biomass (mg).

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 106.8 42.8 85.3 20.8 26.4 12.0 8.4 5.1 Graz. Typical n/a 72.2 16.4 34.5 14.4 10.4 6.0 4.5 2.4 Grass Cut Rest. Plough 17.9 9.3 8.6 5.9 2.6 1.5 1.4 1.7 Cut Rest. Min. Cult. 37.5 13.6 14.0 8.5 7.6 2.4 4.0 3.2 Cut Typical. Plough 11.0 8.7 12.5 4.4 7.4 2.6 4.6 1.4 Cut Typical. Min. Cult. 38.4 6.3 9.8 1.2 26.7 1.2 3.3 0.4 Graz. Rest. Plough 23.6 37.6 22.0 2.6 6.8 20.7 9.5 0.7 Graz. Rest. Min. Cult. 34.0 29.3 28.4 14.5 10.9 5.1 15.7 5.0 Graz. Typical. Plough 7.5 4.3 12.5 5.6 1.8 1.3 5.3 1.2 Graz. Typical. Min. Cult. 13.2 4.1 5.3 6.0 5.4 1.1 1.0 2.6 GL Cut Rest. Plough 131.7 81.3 85.4 41.9 11.9 15.2 11.0 5.0 Cut Rest. Min. Cult. 235.9 86.6 132.6 72.7 35.1 6.8 22.8 1.8 Cut Typical. Plough 74.0 60.6 122.1 59.9 11.2 10.6 21.7 11.0 Cut Typical. Min. Cult. 89.3 51.2 121.5 104.2 31.3 10.6 17.6 21.5 Graz. Rest. Plough 136.6 121.4 53.3 23.2 12.8 9.1 15.4 6.6 Graz. Rest. Min. Cult. 192.8 82.7 75.3 32.8 27.1 10.5 24.8 4.8 Graz. Typical. Plough 65.4 21.4 19.3 26.3 8.2 5.3 1.6 8.7 Graz. Typical. Min. Cult. 74.6 20.5 41.0 23.3 8.1 1.8 3.7 3.8 GLF Cut Rest. Plough 154.3 76.4 61.8 35.8 5.0 13.7 9.0 6.8 Cut Rest. Min. Cult. 340.5 95.4 153.2 42.1 32.8 5.7 29.7 2.9 Cut Typical. Plough 105.1 74.7 143.6 44.5 16.7 3.0 33.9 13.3 Cut Typical. Min. Cult. 83.8 63.2 172.6 68.4 8.7 14.2 32.0 11.4 Graz. Rest. Plough 155.1 162.8 45.3 14.7 4.2 26.6 4.8 3.9 Graz. Rest. Min. Cult. 189.9 109.7 41.0 19.8 21.5 17.4 5.8 6.3 Graz. Typical. Plough 73.0 24.2 24.8 59.2 19.5 4.3 2.1 28.2 Graz. Typical. Min. Cult. 103.9 21.2 107.3 39.2 5.4 4.5 58.9 10.1

190

Appendix 2i: Jealott’s Hill beetle and pollinator abundance.

Beetle abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 232.8 84.3 142.3 42.0 75.5 27.8 26.6 7.8 Graz. Typical n/a 88.8 17.0 42.8 23.3 12.9 4.9 4.5 1.8 Grass Cut Rest. Plough 22.8 15.0 13.5 10.0 5.0 3.0 1.3 2.1 Cut Rest. Min. Cult. 26.3 16.3 19.5 10.8 3.0 1.0 3.1 1.9 Cut Typical. Plough 8.8 15.3 14.5 7.8 4.4 3.8 2.7 2.4 Cut Typical. Min. Cult. 8.5 10.0 11.0 3.0 1.2 1.5 1.6 0.9 Graz. Rest. Plough 24.8 18.3 20.0 7.3 2.3 4.5 2.2 1.7 Graz. Rest. Min. Cult. 19.5 22.3 22.0 14.3 3.3 3.7 9.7 9.6 Graz. Typical. Plough 7.8 5.5 12.0 10.3 0.8 2.5 1.8 2.8 Graz. Typical. Min. Cult. 6.8 5.5 9.8 5.8 3.1 1.7 1.9 1.3 GL Cut Rest. Plough 228.0 116.0 111.0 61.5 35.9 26.6 15.2 10.7 Cut Rest. Min. Cult. 438.0 109.5 182.5 67.3 107.8 10.3 10.2 11.8 Cut Typical. Plough 125.5 95.8 154.5 68.3 17.4 12.7 16.8 14.9 Cut Typical. Min. Cult. 172.8 82.0 184.3 93.3 74.8 17.7 23.4 15.7 Graz. Rest. Plough 161.3 158.8 49.5 28.5 8.1 13.4 4.9 4.3 Graz. Rest. Min. Cult. 245.3 120.5 77.0 36.8 44.9 17.7 5.0 6.9 Graz. Typical. Plough 75.5 28.5 36.8 30.5 14.7 5.5 2.7 7.2 Graz. Typical. Min. Cult. 97.5 27.0 59.5 36.0 17.0 5.2 7.2 6.6 GLF Cut Rest. Plough 242.8 94.3 73.0 39.3 14.9 14.0 6.9 3.9 Cut Rest. Min. Cult. 665.0 106.5 165.8 60.0 88.3 21.0 35.9 5.8 Cut Typical. Plough 191.3 102.0 163.3 57.5 29.8 10.5 19.6 15.6 Cut Typical. Min. Cult. 172.8 93.3 170.8 92.5 22.0 23.6 14.5 11.0 Graz. Rest. Plough 197.3 168.0 34.0 22.5 9.7 32.0 3.2 2.5 Graz. Rest. Min. Cult. 273.5 133.8 38.8 26.8 30.5 25.0 8.9 7.7 Graz. Typical. Plough 94.0 27.8 34.0 32.5 18.3 4.3 3.7 9.8 Graz. Typical. Min. Cult. 128.0 26.8 50.3 46.3 19.2 5.2 6.9 7.8

Total pollinator abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 9.8 8.3 13.5 1.8 1.9 4.3 3.5 0.8 Graz. Typical n/a 10.5 4.3 16.0 4.0 3.8 0.9 2.0 2.7 Grass Cut Rest. Plough 0.3 0.3 2.8 1.8 0.3 0.3 1.0 0.8 Cut Rest. Min. Cult. 0.8 0.0 2.8 0.8 0.5 0.0 0.6 0.3 Cut Typical. Plough 0.5 1.5 1.0 0.5 0.3 1.2 0.4 0.3 Cut Typical. Min. Cult. 1.0 0.5 1.8 2.0 1.0 0.5 0.6 0.8 Graz. Rest. Plough 1.0 0.3 2.8 3.0 0.7 0.3 1.1 1.3 Graz. Rest. Min. Cult. 0.8 0.8 5.5 3.8 0.5 0.3 1.7 1.9 Graz. Typical. Plough 2.0 0.0 2.5 3.3 1.2 0.0 0.6 0.8 Graz. Typical. Min. Cult. 0.8 0.8 2.3 2.3 0.5 0.8 1.6 0.8 GL Cut Rest. Plough 97.8 11.5 43.5 6.8 17.6 2.8 8.0 2.6 Cut Rest. Min. Cult. 85.3 14.3 35.0 7.0 15.4 3.9 3.7 2.5 Cut Typical. Plough 53.5 11.0 28.3 4.5 9.1 1.6 7.6 1.9 Cut Typical. Min. Cult. 49.5 14.3 24.0 7.0 7.0 2.3 2.7 2.3 Graz. Rest. Plough 34.5 27.0 43.0 8.3 19.4 8.5 22.4 3.0 Graz. Rest. Min. Cult. 25.8 18.8 24.5 5.5 13.8 4.2 12.3 1.3 Graz. Typical. Plough 16.3 3.5 9.3 4.0 6.6 1.6 1.3 0.9 Graz. Typical. Min. Cult. 15.8 7.8 15.0 4.8 5.1 3.4 2.4 1.1 GLF Cut Rest. Plough 106.3 15.5 73.0 66.5 18.0 2.3 2.1 11.4 Cut Rest. Min. Cult. 90.8 13.3 81.0 62.8 20.5 2.2 2.9 10.0 Cut Typical. Plough 66.5 16.8 45.0 76.3 6.0 5.5 3.3 6.4 Cut Typical. Min. Cult. 61.8 11.5 57.5 62.8 12.7 3.4 6.4 3.4 Graz. Rest. Plough 41.5 26.3 48.5 35.3 18.5 6.3 7.4 4.9 Graz. Rest. Min. Cult. 37.8 19.5 47.5 40.3 22.1 2.9 8.1 7.7 Graz. Typical. Plough 17.3 4.3 30.0 27.5 5.8 2.7 1.7 2.9 Graz. Typical. Min. Cult. 16.5 2.8 14.0 21.3 3.8 1.1 4.1 2.6

191

Honeybee abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 2.5 1.8 10.5 0.0 1.9 0.8 2.1 0.0 Graz. Typical n/a 0.5 2.3 6.0 2.5 0.3 0.8 1.3 2.5 Grass Cut Rest. Plough 0.0 0.0 1.3 0.0 0.0 0.0 0.8 0.0 Cut Rest. Min. Cult. 0.0 0.0 0.3 0.0 0.0 0.0 0.3 0.0 Cut Typical. Plough 0.0 0.0 0.3 0.3 0.0 0.0 0.3 0.3 Cut Typical. Min. Cult. 0.0 0.0 1.3 0.0 0.0 0.0 0.6 0.0 Graz. Rest. Plough 0.0 0.0 0.3 0.0 0.0 0.0 0.3 0.0 Graz. Rest. Min. Cult. 0.0 0.0 0.3 0.0 0.0 0.0 0.3 0.0 Graz. Typical. Plough 0.0 0.0 0.5 0.5 0.0 0.0 0.5 0.5 Graz. Typical. Min. Cult. 0.0 0.5 0.3 0.3 0.0 0.5 0.3 0.3 GL Cut Rest. Plough 25.8 1.5 18.3 1.0 5.4 0.9 5.2 1.0 Cut Rest. Min. Cult. 22.5 2.8 10.0 0.3 6.3 1.1 3.9 0.3 Cut Typical. Plough 13.5 2.3 9.8 0.5 4.9 0.8 6.6 0.5 Cut Typical. Min. Cult. 11.0 2.0 6.5 1.3 2.6 0.7 1.0 0.5 Graz. Rest. Plough 11.5 3.8 21.5 0.5 6.9 1.4 9.3 0.3 Graz. Rest. Min. Cult. 4.8 4.0 11.5 0.3 3.2 1.5 5.6 0.3 Graz. Typical. Plough 5.5 0.8 2.3 0.5 4.2 0.3 0.6 0.5 Graz. Typical. Min. Cult. 2.3 0.8 2.3 0.3 0.9 0.5 1.1 0.3 GLF Cut Rest. Plough 25.3 1.8 26.8 21.5 5.8 0.6 4.6 6.4 Cut Rest. Min. Cult. 14.3 4.3 27.8 23.5 5.8 0.5 2.9 5.3 Cut Typical. Plough 13.3 3.5 15.8 26.8 2.8 1.8 4.0 1.7 Cut Typical. Min. Cult. 17.3 2.8 19.0 21.8 7.0 1.4 3.4 4.5 Graz. Rest. Plough 10.0 2.5 9.3 8.8 5.7 0.3 1.9 0.8 Graz. Rest. Min. Cult. 8.5 5.3 10.8 10.5 6.8 1.5 2.1 2.4 Graz. Typical. Plough 2.8 0.5 8.3 6.3 1.5 0.3 4.1 1.3 Graz. Typical. Min. Cult. 2.8 0.5 7.8 4.8 1.3 0.3 3.9 1.4

Bumblebee abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 6.8 5.0 1.5 0.0 1.1 3.0 0.6 0.0 Graz. Typical n/a 9.0 1.8 7.8 0.5 3.3 0.8 1.4 0.5 Grass Cut Rest. Plough 0.0 0.0 0.5 0.3 0.0 0.0 0.5 0.3 Cut Rest. Min. Cult. 0.0 0.0 0.5 0.0 0.0 0.0 0.5 0.0 Cut Typical. Plough 0.3 1.0 0.0 0.0 0.3 0.7 0.0 0.0 Cut Typical. Min. Cult. 1.0 0.0 0.0 0.3 1.0 0.0 0.0 0.3 Graz. Rest. Plough 0.0 0.0 1.0 0.5 0.0 0.0 0.7 0.3 Graz. Rest. Min. Cult. 0.3 0.3 1.8 2.0 0.3 0.3 1.0 1.1 Graz. Typical. Plough 1.3 0.0 1.0 1.0 1.3 0.0 1.0 0.7 Graz. Typical. Min. Cult. 0.0 0.3 0.3 0.3 0.0 0.3 0.3 0.3 GL Cut Rest. Plough 53.0 7.8 17.8 3.0 12.5 1.1 0.8 2.0 Cut Rest. Min. Cult. 41.8 10.3 21.3 3.8 3.5 3.9 1.9 1.0 Cut Typical. Plough 33.3 7.3 13.0 2.3 4.2 1.7 4.5 1.6 Cut Typical. Min. Cult. 30.3 10.5 11.8 3.5 5.1 1.8 2.7 1.3 Graz. Rest. Plough 16.5 22.3 15.3 4.5 10.7 8.0 10.9 1.8 Graz. Rest. Min. Cult. 17.5 13.0 10.5 2.5 9.7 3.5 5.9 0.3 Graz. Typical. Plough 8.5 2.0 3.0 0.5 3.1 1.1 1.3 0.3 Graz. Typical. Min. Cult. 11.8 6.0 6.8 2.3 4.2 2.4 1.6 0.9 GLF Cut Rest. Plough 57.0 7.0 25.8 18.5 7.8 1.1 4.2 2.7 Cut Rest. Min. Cult. 58.8 6.5 30.0 14.3 8.7 1.4 3.4 2.6 Cut Typical. Plough 42.0 10.3 21.5 17.8 2.7 2.5 2.6 3.4 Cut Typical. Min. Cult. 36.0 6.5 29.5 16.8 4.4 1.8 3.1 3.0 Graz. Rest. Plough 19.0 16.3 18.0 10.8 7.5 6.1 0.9 4.5 Graz. Rest. Min. Cult. 22.5 10.5 14.8 11.8 9.9 1.7 2.3 2.8 Graz. Typical. Plough 10.3 3.5 4.0 8.8 4.0 2.5 1.5 1.1 Graz. Typical. Min. Cult. 12.8 2.3 4.3 6.0 3.4 0.9 0.6 0.9

192

Butterfly abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.5 1.5 0.5 1.8 0.3 0.9 0.3 0.8 Graz. Typical n/a 0.5 0.3 0.8 1.0 0.3 0.3 0.5 0.4 Grass Cut Rest. Plough 0.3 0.0 0.8 1.5 0.3 0.0 0.5 0.6 Cut Rest. Min. Cult. 0.8 0.0 1.8 0.8 0.5 0.0 0.6 0.3 Cut Typical. Plough 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3 Cut Typical. Min. Cult. 0.0 0.5 0.3 1.8 0.0 0.5 0.3 0.6 Graz. Rest. Plough 1.0 0.3 1.5 2.3 0.7 0.3 0.6 1.0 Graz. Rest. Min. Cult. 0.5 0.5 3.0 1.8 0.3 0.3 0.8 1.0 Graz. Typical. Plough 0.8 0.0 0.5 1.3 0.8 0.0 0.3 0.5 Graz. Typical. Min. Cult. 0.8 0.0 1.3 1.5 0.5 0.0 0.9 0.6 GL Cut Rest. Plough 9.0 2.3 2.8 1.5 2.3 0.9 0.9 0.6 Cut Rest. Min. Cult. 9.0 1.3 2.0 1.8 0.9 0.9 1.1 0.9 Cut Typical. Plough 3.0 1.5 3.0 1.3 2.4 0.5 0.7 0.3 Cut Typical. Min. Cult. 4.3 1.5 1.3 2.3 1.9 0.6 0.9 0.8 Graz. Rest. Plough 2.8 0.3 3.3 1.8 0.5 0.3 1.3 0.6 Graz. Rest. Min. Cult. 2.3 1.3 1.3 1.8 0.9 0.5 0.8 0.9 Graz. Typical. Plough 1.3 0.5 1.5 3.0 0.3 0.3 1.0 0.8 Graz. Typical. Min. Cult. 1.5 0.3 2.0 1.8 0.6 0.3 0.4 0.8 GLF Cut Rest. Plough 7.5 1.3 2.8 3.5 2.5 0.5 1.0 0.9 Cut Rest. Min. Cult. 6.0 0.8 1.8 3.5 3.2 0.5 1.1 0.5 Cut Typical. Plough 4.8 2.0 1.0 4.0 2.2 1.0 0.4 1.2 Cut Typical. Min. Cult. 3.0 0.8 2.3 3.3 0.7 0.5 1.7 1.0 Graz. Rest. Plough 3.5 4.5 2.5 2.8 1.2 0.3 1.0 0.3 Graz. Rest. Min. Cult. 2.5 2.0 0.8 3.0 1.3 0.4 0.5 1.2 Graz. Typical. Plough 0.8 0.0 4.5 2.8 0.5 0.0 2.2 0.5 Graz. Typical. Min. Cult. 0.8 0.0 1.0 3.0 0.5 0.0 0.4 0.4

Hoverfly abundance

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.0 0.0 1.0 0.0 0.0 0.0 1.0 0.0 Graz. Typical n/a 0.5 0.0 1.5 0.0 0.5 0.0 1.2 0.0 Grass Cut Rest. Plough 0.0 0.3 0.3 0.0 0.0 0.3 0.3 0.0 Cut Rest. Min. Cult. 0.0 0.0 0.3 0.0 0.0 0.0 0.3 0.0 Cut Typical. Plough 0.0 0.3 0.5 0.0 0.0 0.3 0.3 0.0 Cut Typical. Min. Cult. 0.0 0.0 0.3 0.0 0.0 0.0 0.3 0.0 Graz. Rest. Plough 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.3 Graz. Rest. Min. Cult. 0.0 0.0 0.5 0.0 0.0 0.0 0.5 0.0 Graz. Typical. Plough 0.0 0.0 0.5 0.5 0.0 0.0 0.3 0.5 Graz. Typical. Min. Cult. 0.0 0.0 0.5 0.0 0.0 0.0 0.5 0.0 GL Cut Rest. Plough 10.0 0.0 4.8 1.3 5.8 0.0 1.8 0.9 Cut Rest. Min. Cult. 12.0 0.0 1.5 1.3 5.4 0.0 0.9 1.3 Cut Typical. Plough 3.8 0.0 2.5 0.5 2.1 0.0 0.9 0.5 Cut Typical. Min. Cult. 3.8 0.3 4.0 0.0 1.6 0.3 1.1 0.0 Graz. Rest. Plough 3.8 0.8 3.0 1.5 1.9 0.3 1.2 0.5 Graz. Rest. Min. Cult. 1.3 0.5 1.3 0.8 1.3 0.5 0.5 0.5 Graz. Typical. Plough 1.0 0.0 2.5 0.0 0.6 0.0 0.9 0.0 Graz. Typical. Min. Cult. 0.3 0.8 4.0 0.5 0.3 0.8 1.5 0.3 GLF Cut Rest. Plough 16.3 5.5 17.5 22.8 5.8 1.2 1.7 2.8 Cut Rest. Min. Cult. 11.5 1.8 21.5 21.0 5.9 0.3 1.3 3.1 Cut Typical. Plough 6.5 0.5 6.3 26.3 2.4 0.5 1.1 5.8 Cut Typical. Min. Cult. 5.5 1.0 6.5 20.5 1.9 0.7 1.3 3.2 Graz. Rest. Plough 8.5 2.8 18.8 13.0 4.1 1.6 4.2 0.9 Graz. Rest. Min. Cult. 4.3 1.8 21.3 15.0 4.3 1.2 5.3 3.1 Graz. Typical. Plough 3.5 0.3 11.8 9.8 2.0 0.3 5.7 1.2 Graz. Typical. Min. Cult. 0.3 0.0 0.8 7.5 0.3 0.0 0.3 2.0

193

Appendix 2j: Jealott’s Hill beetles and pollinator species richness.

Beetle species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 11.8 10.0 13.5 8.3 0.8 1.5 0.9 0.8 Graz. Typical n/a 8.8 6.8 12.0 7.3 0.5 1.2 1.4 1.3 Grass Cut Rest. Plough 10.5 6.3 6.5 3.8 1.6 0.5 1.6 0.5 Cut Rest. Min. Cult. 13.8 7.5 6.5 5.0 2.0 0.9 1.2 0.7 Cut Typical. Plough 5.8 5.5 6.0 4.3 2.3 0.9 1.1 0.9 Cut Typical. Min. Cult. 5.0 4.5 5.8 2.5 0.9 0.5 0.8 0.6 Graz. Rest. Plough 11.8 8.0 6.3 3.0 0.9 2.0 0.9 0.4 Graz. Rest. Min. Cult. 11.8 8.5 7.5 4.8 1.7 0.9 2.5 1.5 Graz. Typical. Plough 5.3 4.5 6.8 4.8 1.1 1.8 0.3 0.9 Graz. Typical. Min. Cult. 5.5 4.3 5.5 4.0 1.9 1.1 0.5 0.9 GL Cut Rest. Plough 21.8 17.8 23.3 14.5 3.4 1.8 2.2 1.5 Cut Rest. Min. Cult. 20.8 17.8 20.0 16.3 1.4 1.1 1.4 1.4 Cut Typical. Plough 15.0 14.5 22.5 15.8 1.5 1.6 1.7 1.7 Cut Typical. Min. Cult. 14.0 14.0 19.5 15.8 2.3 2.7 1.3 1.0 Graz. Rest. Plough 19.3 22.3 19.8 12.0 1.4 0.9 1.1 1.6 Graz. Rest. Min. Cult. 21.3 18.3 20.8 14.5 2.7 0.9 1.5 1.0 Graz. Typical. Plough 16.8 9.5 11.3 10.8 1.4 1.3 0.5 2.2 Graz. Typical. Min. Cult. 13.8 8.3 14.8 10.8 1.3 0.5 0.6 0.8 GLF Cut Rest. Plough 23.5 18.8 21.5 13.3 0.9 3.1 0.9 1.3 Cut Rest. Min. Cult. 21.0 18.3 23.5 13.8 2.2 1.3 2.5 1.4 Cut Typical. Plough 16.0 14.5 22.3 13.5 1.8 1.0 2.1 0.9 Cut Typical. Min. Cult. 16.3 15.3 25.0 15.8 1.3 1.2 1.5 1.1 Graz. Rest. Plough 22.5 24.5 14.5 9.5 1.4 2.2 0.6 0.9 Graz. Rest. Min. Cult. 21.8 19.8 16.0 11.0 1.7 1.8 2.3 2.8 Graz. Typical. Plough 19.3 11.3 13.3 8.8 2.4 0.6 1.4 0.5 Graz. Typical. Min. Cult. 14.3 8.5 16.0 11.5 1.9 1.0 1.5 1.7

Bee (Bombus and Apis mellifera) species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 2.8 1.8 2.3 0.0 0.9 0.5 0.5 0.0 Graz. Typical n/a 3.0 1.8 3.0 0.8 1.0 0.3 0.4 0.5 Grass Cut Rest. Plough 0.0 0.0 0.8 0.3 0.0 0.0 0.5 0.3 Cut Rest. Min. Cult. 0.0 0.0 0.5 0.0 0.0 0.0 0.3 0.0 Cut Typical. Plough 0.3 0.5 0.3 0.3 0.3 0.3 0.3 0.3 Cut Typical. Min. Cult. 0.5 0.0 0.8 0.3 0.5 0.0 0.3 0.3 Graz. Rest. Plough 0.0 0.0 1.0 0.5 0.0 0.0 0.4 0.3 Graz. Rest. Min. Cult. 0.3 0.3 1.0 1.0 0.3 0.3 0.6 0.4 Graz. Typical. Plough 0.8 0.0 0.8 1.0 0.8 0.0 0.5 0.6 Graz. Typical. Min. Cult. 0.0 0.5 0.5 0.5 0.0 0.5 0.3 0.3 GL Cut Rest. Plough 5.8 3.0 4.3 1.5 0.3 0.6 0.5 0.6 Cut Rest. Min. Cult. 5.5 3.0 4.3 2.3 0.3 0.7 0.5 0.5 Cut Typical. Plough 4.8 2.8 3.3 1.5 0.3 0.3 0.8 0.6 Cut Typical. Min. Cult. 5.3 3.0 4.0 2.5 0.5 0.0 0.4 0.9 Graz. Rest. Plough 3.8 4.3 2.8 1.8 0.8 0.5 0.8 0.6 Graz. Rest. Min. Cult. 4.5 3.3 3.0 2.0 0.5 0.8 0.4 0.4 Graz. Typical. Plough 3.8 1.8 2.0 0.8 0.8 0.3 0.4 0.5 Graz. Typical. Min. Cult. 4.3 2.5 3.0 1.8 1.1 0.6 0.4 0.6 GLF Cut Rest. Plough 5.3 3.0 4.5 3.8 0.3 0.4 0.3 0.3 Cut Rest. Min. Cult. 5.8 3.0 3.8 3.8 0.3 0.4 0.6 0.3 Cut Typical. Plough 5.3 2.8 4.8 3.8 0.3 0.5 0.3 0.5 Cut Typical. Min. Cult. 5.8 2.8 4.5 4.3 0.3 0.5 0.5 0.2 Graz. Rest. Plough 4.0 3.3 3.0 3.5 0.8 0.5 0.0 0.3 Graz. Rest. Min. Cult. 4.8 3.8 3.5 3.8 0.6 0.3 0.3 0.3 Graz. Typical. Plough 3.8 1.3 3.0 4.0 0.9 0.5 0.4 0.0 Graz. Typical. Min. Cult. 4.5 1.8 3.0 3.3 0.6 0.5 0.4 0.2

194

Butterfly species richness

Seed mix Man. Timing Cult. Means SE 2009 2010 2011 2012 2009 2010 2011 2012 Original grassland Cut Typical n/a 0.5 0.8 0.5 1.0 0.3 0.3 0.3 0.4 Graz. Typical n/a 0.5 0.3 0.5 0.8 0.3 0.3 0.3 0.3 Grass Cut Rest. Plough 0.3 0.0 0.8 0.8 0.3 0.0 0.5 0.3 Cut Rest. Min. Cult. 0.8 0.0 1.3 0.8 0.5 0.0 0.5 0.3 Cut Typical. Plough 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3 Cut Typical. Min. Cult. 0.0 0.3 0.3 0.8 0.0 0.3 0.3 0.3 Graz. Rest. Plough 1.0 0.3 1.3 1.3 0.7 0.3 0.5 0.5 Graz. Rest. Min. Cult. 0.5 0.5 1.5 1.0 0.3 0.3 0.3 0.7 Graz. Typical. Plough 0.8 0.0 0.5 0.8 0.8 0.0 0.3 0.3 Graz. Typical. Min. Cult. 0.8 0.0 0.8 0.8 0.5 0.0 0.5 0.3 GL Cut Rest. Plough 2.5 1.5 1.5 1.0 0.5 0.3 0.3 0.4 Cut Rest. Min. Cult. 2.8 0.8 1.3 1.0 0.3 0.5 0.5 0.4 Cut Typical. Plough 0.8 0.8 2.0 1.0 0.5 0.3 0.7 0.0 Cut Typical. Min. Cult. 1.8 0.8 0.8 1.0 0.9 0.3 0.5 0.0 Graz. Rest. Plough 1.5 0.3 1.8 1.0 0.3 0.3 0.5 0.4 Graz. Rest. Min. Cult. 1.0 1.3 0.8 1.0 0.4 0.5 0.5 0.4 Graz. Typical. Plough 1.3 0.5 0.8 1.0 0.3 0.3 0.5 0.0 Graz. Typical. Min. Cult. 0.8 0.3 1.3 0.8 0.3 0.3 0.3 0.3 GLF Cut Rest. Plough 2.0 0.8 1.3 1.5 0.6 0.3 0.3 0.3 Cut Rest. Min. Cult. 1.8 0.5 1.0 1.5 0.6 0.3 0.4 0.3 Cut Typical. Plough 1.3 1.3 0.8 1.5 0.5 0.3 0.3 0.3 Cut Typical. Min. Cult. 1.5 0.5 1.3 1.0 0.3 0.3 0.8 0.0 Graz. Rest. Plough 2.0 1.3 1.0 1.0 0.0 0.3 0.4 0.0 Graz. Rest. Min. Cult. 1.5 1.5 0.8 1.3 0.5 0.5 0.5 0.3 Graz. Typical. Plough 0.8 0.0 1.0 1.5 0.5 0.0 0.4 0.3 Graz. Typical. Min. Cult. 0.8 0.0 0.8 1.3 0.5 0.0 0.3 0.3

195

Appendix 3: North Wyke soil descriptions of unsown ‘existing grassland’ control plots.

Summary

At North Wyke there were eight unsown ‘existing grassland’ control plots. Four of these control plots (18, 22, 47 and 62) were typically grazed, comprising of continuous grazing with cattle from April/May to October. The other four plots (2, 32, 38 and 70) were managed by a typical cutting regime achieved by a first cut in May/June and a second cut in August/September.

On the 8 existing sward, typically managed plots, soil structural degradation was most often classed as moderate, based on the presence of weakly developed structure in the topsoil and immediate subsoil overlying clay, or moderate poor, where platiness occurs, probably resulting from past compaction. There is one example of low degradation (plot 47, grazed), where moderately developed structure was found.

Introduction and Methodology

There is an increasing amount of evidence to indicate that modern agriculture is causing serious degradation of soil structure in both arable and grassland systems (eg Palmer et al 2008). This examination of soil structure on the WEB control plots at Rowden uses an approach developed in recent years in work funded by the Environment Agency to investigate the nature and degree of soil structural degradation and its contribution to run-off, flooding and pollution of water bodies, and to determine the success or otherwise of mitigation strategies.

This work was undertaken on 24th and 27th March 2009. Weather was cool and showery, and ground conditions were generally moist. Plots selected for this investigation (2, 18, 22, 32, 38, 47, 62 and 70) had not been cultivated/reseeded or managed by minimum tillage for about 10 years. Soil profile descriptions were made at an equivalent location in each plot, measuring from the south west corner 5 paces to the east, along the southern boundary, then 5 paces north. After descriptions were made, each location was marked by a blue post.

At each sample point, a small pit about 30 x 30 cm was excavated with a spade down to about 40 to 50 cm depth. A Dutch auger was then used in the base of the pit to obtain samples of the soil profile down to a maximum of 120 cm depth, or less if the auger was stopped by bedrock or hard stony material. Soil profile characteristics were described using the terminology of Hodgson (1997).

An updated map and description of the soils of North Wyke and Rowden has been produced by Harrod and Hogan (2008), and can be found on the North Wyke web site. The elevated area known as Enclosures has been mapped almost entirely as Halstow soils. These are non-calcareous pelosols (Avery 1980) defined as comprising clayey material over lithoskeletal mudstone, shale or slate (Clayden and Hollis (1984), ie clay soils over stony material within 80 cm depth. Where soil profiles differ from the Halstow series, they are indicated here as a variant as defined by their differing characteristics. Some of these profiles could be defined as different series, though these are not specified here to avoid unnecessary complication. A small area of wetter clayey pelo-stagnogley soils of the Hallsworth series is shown on lower slopes in the south-west part of Enclosures, and forms part of a much more extensive area of these soils found across the lower ground of Rowden.

The assessment of soil structural degradation is based on the scheme developed by Palmer et al (2006) with some more recent modifications (Palmer pers com). The main features are summarised in Table 1. The scheme has been used as a predictor of enhanced run-off and to identify the needs for remedial actions to reduce the risks of flooding and the pollution of water bodies. In this survey, the evidence for structural degradation focuses mainly on the characteristics of the upper part of the soil profile, in particular the soil structure and moisture of the topsoil and immediate subsoil horizons. In every case, clay subsoils occur at variable depth, and moisture tends to decline in these deeper layers, typical of slowly impermeable soils not affected by shallow groundwater. No evidence of poaching or wheelings is not found on these experimental plots.

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Table 1 Classes of soil structural degradation

Degradation Description of hydrological Management Soil degradation features class implications system Severe (S) Soil degradation generates All systems: Extensive rill erosion on slopes, sufficient enhanced runoff to arable and depositional fans on footslopes and level cause widespread erosion that grassland ground. Plus most characteristics of High is not confined to wheelings degradation

High (H) Soil degradation generates Arable Slaked or capped topsoil; wheelings 5 cm enhanced runoff across whole or more in depth; structural change in fields where slopes allow topsoil or immediate subsurface layer; compaction; changes to vertical wetness gradient; local erosion in wheelings

Grassland Extensively poached surface or wheelings 5 cm or deeper; damage to topsoil or immediate subsurface structure (apedal or weak coarse angular blocky structure); changes to vertical wetness gradient

Moderate (M) Soil degradation generates Arable Slaked or partly slaked topsoil; wheelings localised areas of enhanced generally less than 5 cm deep; topsoil runoff where slopes allow structural change

Grassland Slight poaching (locally severe); weak subsurface structure/compaction; where structural degradation requires urgent amelioration, but damage does not qualify for High, class Moderate Poor (MP) has been used

Low (L) Insignificant enhanced runoff All systems: Few signs of enhanced runoff generation arable and mechanisms present, but can show signs grassland of localised poaching and standing water as long as the whole profile maintains a good soil structure

Results

Soil profile descriptions

Plot 2: Typical cutting Soil profile class: Halstow map unit; Halstow variant, loamy over clayey Vegetation: permanent grass with rush Slope: 2 degrees convex

Depth (cm) Description Horizon 0-20 Dark brown to brown (10YR 4/3) silty clay loam with common fine rusty mottles; Ah few small sandstones; moist; weak, locally moderate, fine angular blocky structure, with some coarse platy structure developed; moderately firm soil strength; sharp even boundary (probably base of old plough layer)

197

20-40 Very pale brown (10YR 7/4) silty clay loam with many medium mottles of Bw(g) yellowish red (5YR 4/6) and many medium manganese coats; common medium sandstones; moist; moderate very coarse prismatic structure; moderately strong soil strength

40-50 Very pale brown (10YR 7/3) silty clay with many medium mottles of yellowish red Bg (5YR 5/6); slightly moist; over hard stone

Soil structural degradation class: moderate Features: some poor structural development in the topsoil including platiness

Plot 18: Typical grazing Soil profile class: Halstow map unit; Halstow series Vegetation: permanent grass Slope: 4 degrees convex

Depth (cm) Description Horizon 0-9 Dark greyish brown (10YR 4/2) silty clay loam with many fine rusty mottles; Ah common small sandstones; moist; moderate fine angular blocky structure; moderately firm soil strength (some strength imparted by roots mat); abundant (many below 5 cm) very fine fibrous roots in mat

9-20 Brown (10YR 5/3) silty clay loam with common (many below 16 cm) rusty mottles; Ah(g) common small sandstones, and a concentration of large sandstones at the base of the horizon; moist; very weak angular blocky structure; moderately firm soil strength; common very fine fibrous roots, many dead in fissures; sharp even boundary (base of old plough layer?)

20-45 Very pale brown (10YR 7/4) silty clay with many medium mottles of strong brown Bw(g) (7.5YR 5/6) and light brownish grey (10YR 6/2) on ped faces; few small stones; moist; moderate very coarse prismatic structure; moderately strong soil strength

45-65 Light grey (N7/-) silty clay with many medium mottles of strong brown (7.5YR Bg 5/8); moist; over hard stone

Soil structural degradation class: moderate Features: weakly structured below the topsoil

Plot 22: Typical grazing Soil profile class: Halstow map unit; Halstow series Vegetation: permanent grass Slope: 4 degrees convex

Depth (cm) Description Horizon 0-9 Dark brown to brown (10YR 4/3) silty clay loam with common fine rusty mottles; Ah(g)1 common small sandstones; moist; moderate fine and medium granular structure; moderately firm soil strength; many very fine fibrous roots

9-21 Brown (10YR 5/3) silty clay loam with common (many below 16 cm) rusty mottles; Ah(g)2 common small sandstones; moist; weak fine angular blocky structure; moderately

198

firm soil strength; common very fine fibrous roots

21-40 Very pale brown (10YR 7/3) clay with many medium mottles of yellowish brown Bw(g) (10YR 5/6) and pale brown (10YR 6/3) on ped faces; few small sandstones (abundant in top 5 cm of horizon); moist; weak very coarse prismatic structure; very firm soil strength

40-55 Light grey (N7/-) clay with many medium mottles of strong brown (7.5YR 5/8); Bg slightly moist; over hard stone

Soil structural degradation class: moderate Features: weakly structured below the topsoil

Plot 32: Typical cutting Soil profile class: Halstow map unit; Halstow series Vegetation: permanent grass Slope: 4 degrees convex

Depth (cm) Description Horizon 0-34 Dark brown to brown (10YR 4/3) silty clay loam; few small and medium Ah sandstones; moist; weak fine and medium granular structure 0-7 cm, otherwise very weak fine angular blocky structure, and some weak coarse platiness; moderately firm soil strength; many very fine fibrous roots

34-50 Yellow (10YR 7/6) silty clay loam to silty clay with common fine and medium faint Bw(g) mottles of yellowish brown (10YR 5/6); many large sandstones; moist; massive; very firm soil strength; auger stopped on hard stone

Soil structural degradation class: moderate poor Features: deep brown but weakly structured topsoil over compact massive subsoil

Plot 38: Typical cutting Soil profile class: Halstow map unit; Halstow stagnogley marginal to loamy variant Vegetation: permanent grass Slope: 4 degrees convex

Depth (cm) Description Horizon 0-6 Dark greyish brown to brown (10YR 4/2 to 5/3) silty clay loam with many fine Ah rusty mottles; common small and medium sandstones; moist; weak fine and medium granular structure; moderately firm soil strength; abundant fine fibrous roots; earthworms present

6-35 Greyish brown (10YR 5/2) silty clay loam to silty clay; common medium and large Bw(g) sandstones; moist; weak coarse platy structure; moderately firm soil strength; common fine fibrous roots

35-45 Grey (5Y 5/1) clay loam to clay with common fine faint mottles of yellowish brown Bg (10YR 5/6); few small sandstones; moist; weak medium to coarse prismatic structure; very firm soil strength; common fine fibrous roots

199

45-120 Light grey (N7/-) silty clay with many medium and coarse mottles of strong brown BCg (7.5YR 5/8); common stones, increasing with depth; moist, becoming slightly moist and with bluer colour below 90 cm depth; over hard stone

120+ Wetter material too stony to auger and sound of venting gas (presumably in more Cg porous material carrying shallow groundwater)

Soil structural degradation class: moderate poor Features: weakly structured platy subsoil immediately below topsoil.

Plot 47: Typical grazing Soil profile class: Halstow map unit; Halstow stagnogley variant Vegetation: permanent grass Slope: 5 degrees convex

Depth (cm) Description Horizon 0-8 Brown (10YR 4-5/3) silty clay loam with common fine rusty mottles; common Ah small sandstones; moist; moderate fine and medium granular structure; moderately firm soil strength; abundant fine fibrous roots; earthworms present

8-27 Brown (10YR 5/3) silty clay loam with many fine rusty mottles; common medium Bw and large sandstones; moist; moderate fine angular blocky structure; moderately firm soil strength; many very fine fibrous roots; earthworms present

27-50 Pinkish grey (7.5Y 7/2) clay with many medium mottles of strong brown (7.5YR Bg 5/8); few stones; moist; moderate very coarse prismatic structure; moderately strong soil strength

50-70 Light grey (N7/-) clay with many medium and coarse mottles of strong brown BCg (7.5YR 5/8); slightly moist

70+ Slightly moist shaly material

Soil structural degradation class: low Features: moderately developed structure throughout; gleyed with 30 cm of the surface

Plot 62: Typical grazing Soil profile class: Halstow map unit; Halstow stagnogley variant Vegetation: permanent grass with rush Slope: 5 degrees convex

Depth (cm) Description Horizon 0-8 Dark brown to brown (10YR 4/3) silty clay loam with common fine rusty mottles; Ah common small sandstones; moist; moderate fine and medium granular structure; moderately firm soil strength; many fine fibrous roots; earthworms present

8-23 Brown (10YR 5/3) silty clay loam with common fine and medium mottles of Bw(g) yellowish brown (10YR 5/6); common small sandstones; moist; weak fine angular blocky structure; moderately firm soil strength; common fine fibrous roots

23-50 Pale brown (10Y 6/3) silty clay with many fine and medium mottles of strong Bg brown (7.5YR 5/8) and yellowish red (5YR 5/6), and greyish red faces; common medium sandstones; moist; moderate coarse prismatic structure; moderately strong

200

soil strength; common roots, especially in fissures

50-90 Bluish grey (5B 5/1) clay with many fine and medium mottles of strong brown BCg (7.5YR 5/6); much soft weathered shaly material and hard sandstones; slightly moist; auger stopped on hard stone

Soil structural degradation class: moderate Features: subsoil structure weakly developed; gleyed within 30 cm depth

Plot 70: Typical cutting Soil profile class: Halstow map unit; Halstow series Vegetation: permanent grass with rush Slope: 4 degrees convex

Depth (cm) Description Horizon 0-7 Dark brown to brown (10YR 4/3) silty clay loam with common fine rusty mottles; Ah few small sandstones; very moist; weak fine and medium granular structure; moderately firm soil strength; many fine and very fine fibrous roots; earthworms present

7-28 Brown (10YR 5/3) silty clay loam with common fine mottles of yellowish brown Bw(g)1 (10YR 5/6); common small and medium sandstone and shale fragments; moist; weak very coarse platy structure; very firm soil strength; common fine fibrous roots

28-45 Pale brown (10Y 5/3) clay with many coarse mottles of yellowish brown (10YR Bw(g)2 5/8), and with light brownish grey (10YR 6/2) on ped faces; very few stones; slightly moist; weak very coarse prismatic structure; moderately strong soil strength

45-90 Light grey (N 7/-) clay with many medium mottles of yellowish red (5YR 5/8); BCg moist; auger stopped on hard stone

Soil structural degradation class: moderate poor Features: soil wetness decreases with depth indicating impedance of downward drainage

Summary of findings Among the descriptions of the plots examined here, the main variations from the Halstow concept are in depth to stony drift or bedrock, or in the thickness of superficial loamy (silty clay loam) material overlying the clay. These are described here as variants of the Halstow series, though some would meet the definitions of other soil series. Soil profiles meet the definition of Halstow series in 4 of the 8 plots examined.

In most cases there is a relatively sharp and even boundary between horizons at around 30 cm depth, which is likely to indicate the base of a former plough layer. Above this, there is usually a thin upper horizon often with granular structure resulting from some years of subsequent grassland use, below which the structure is generally coarser and/or less well developed and blocky.

Structural degradation is most often classed as moderate, based on the presence of weakly developed structure, or moderate poor, where platiness occurs. In the one example of low degradation (plot 47, grazed), structural development is moderate. Results are summarised in Table 2.

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Table 2 Summary of results

Plot number Typical Structural degradation Soil series or variant (Existing grassland) management class 2 cut M Variant 18 grazed M Halstow 22 grazed M Halstow 32 cut MP Halstow 38 cut MP Variant 47 grazed L Variant 62 grazed M Variant 70 cut MP Halstow

References Avery,B.W. (1980). Soil Classification for England and Wales (Higher Categories). Soil Surv. Tech. Monogr. No. 14. Clayden, B. and Hollis, J.M. (1984). Criteria for Differentiating Soil Series. Soil Surv. Tech. Monogr. No. 17. Palmer, R.C., Burton, R.G.O., Hannam, J.A. and Creamer, R. (2006). Comparison of soil structural conditions in Tone and Parrett catchments during winter periods 2002-03 and 2005-06. Unpublished final report for JAF Project FWAG (Somerset) and the Environment Agency: NSRI Contract No. YE 200039V. National Soil Resources Institute, Cranfield University.

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Appendix 4: Soil infiltration measurements of unsown ‘existing grassland’ control plots at North Wyke.

Introduction

Land use and management is considered to play an important role in regulating the absorption of rainfall and generation of runoff. Surface runoff is generated when rainfall exceeds the speed at which the soil can adsorb water i.e. the soil’s infiltration capacity. Infiltration is affected by topsoil porosity and water content. A reduction in infiltration capacity is caused by soil structural damage resulting from, for example, raindrop impact on a bare surface, or by smearing and reduction in pore space (specifically larger pores) by compaction resulting from trafficking or stocking under wet conditions when soil bearing strength is reduced. The impacts are expressed on the ground surface as wheelings and poaching. Infiltration is also reduced as the soil wets up due to a reduction in forces moving the water down through the profile.

Method

Infiltration was measured as saturated hydraulic conductivity using a falling head double ring infiltrometer (Scotter et al., 1982, McKenzie et al., 2002). The technique involves using an inner and outer metal ring inserted into the ground. The rings are supplied with a manually controlled head of water, and the saturated infiltration rate (also known as saturated hydraulic conductivity) is determined when vertical flow within the inner ring is at a steady state. Water in the outer ring maintains a saturated zone around the area of measurements (the inner ring), and reduces the risk of lateral flow from below the inner ring affecting the results. The rings used were of 20 and 30 cm diameter. The rings were knocked about 5 cm into the ground, using a soft-headed mallet and a piece of wood to prevent damage. The rings have one bevelled edge to ease insertion into the ground, though where a root mat was developed, it was found advisable also to cut around the base of the ring before knocking it in to the required depth.

Once the rings had been set in the ground, water was poured in and a tape measure used to record the depth of the water level below the top within the inner ring at timed intervals. Initially the water level falls more rapidly as the soil becomes saturated, after which a steady state infiltration rate is reached that indicates saturated hydraulic conductivity has been achieved. The results for individual experimental plots are shown on graphs later in the appendix; rates measured in cm h-1 are indicated in Table 1, using the infiltration categories of Landon (1991) shown in Table 2, together with the structural degradation classes previously described in the soil survey of these plots (Hogan 2009).

Infiltration tests were carried out in pairs, to give an indication of local variability, though this could be further improved by the use of larger diameter rings. Following each infiltration test, the rings were removed and soil profile features recorded, noting how far the downward movement of the wetting front extended, and especially where this may have been halted by the presence of an observable layer of lower permeability and/or poorer/coarser structural development.

Results

The outcomes of the infiltration tests are shown as plots in the Appendix, while Table 1 compares these results with the soil structural degradation assessments, as described by Hogan (2009); for each experimental plot the two replicate infiltration test results are shown. The majority of sites were assigned to the degradation class moderate or moderate poor, with one single assessment of low found in plot 47. Infiltration rates were mostly in the range 0.6 to 10.4 cm h-1 (moderately slow –moderately rapid), a general level of similarity in results which might be expected from soils of the same Halstow map unit (Harrod and Hogan 2008) having a similar history of land management. However, of the extreme values recorded, (the fastest was 15.6 cm h-1 (rapid) in

203 plot 2 and the slowest 0.6 cm h-1 (moderately slow) in plot 47), the replicate measurements differed by two and one infiltration class respectively (Table 2), indicating the localised soil variability found within the experimental plots.

The extreme values reflect the variance of the soil profile from the central concept of the Halstow series (a predominantly clayey soil profile classed as a pelosol). In plot 2, the more rapid infiltration rate was found in what might be regarded as a drier variant (loamy topsoil rather than clayey throughout), while the slowest was in plot 47, a wetter version of Halstow more akin to a pelostagnogley of the Hallsworth series, which most commonly occupies footslopes at North Wyke and Rowden (Harrod and Hogan 2008).

Table 1 Summary of results

Plot Typical Structural Soil series or Infiltration number management degradation variant Rate (cm h-1) Category (existing class grassland) 2 cut M Halstow 15.6 / 2.4 rapid / moderate variant 18 grazed M Halstow Not measured - 22 grazed M Halstow 7.2 / 2.0 moderately rapid / moderately slow - slow 32 cut MP Halstow 9.6 / 10.4 moderately rapid / moderately rapid 38 cut MP Halstow 4.6 / 4.2 moderate / variant moderate 47 grazed L Halstow 4.8 / 0.6 moderate / variant moderately slow 62 grazed M Halstow 2.2 / 0.7 moderate / variant moderately slow 70 cut MP Halstow 1.4 / 2.6 moderately slow / moderate

Table 2 Infiltration categories

Class Infiltration category Infiltration rate (cm h-1) 1 Very slow <0.1 2 Slow 0.1-0.5 3 Moderately slow 0.5-2.0 4 Moderate 2.0-6.0 5 Moderately rapid 6.0-12.5 6 Rapid 12.5-25.0 7 Very rapid >25.0

Discussion

The variations found in infiltration rates measured for individual plots (and consequent soil degradation classes) reflects very localised variations in soil properties affecting infiltration, while at the same time highlighting problems of interpreting soil structural degradation where there is a lack of ground surface features, which comprise key criteria in the methodology described by Palmer et al 2006; in addition the

204 approach is intended to assess whole fields, or at least identifiable areas having discrete management-induced features such as wheelings or poaching.

The work highlights the importance of having data such as infiltration measurements to support soil structural assessments based on observable profile characteristics and ground surface features. This is particularly important in the case of experimental plots such as at Rowden, where land use and management is more regulated for the purposes of research than would usually be the case on most farms, with the result that commonly observed agricultural management-induced features may not be present to provide visual evidence in soil quality assessments.

References

Hogan, D.V. (2009). Soil descriptions of selected WEB plots at Rowden. Unpublished report to North Wyke Research. Landon, J.R. (Ed.) (1991). Booker Tropical Soil Manual. Booker Tate Ltd. Ch 6, p69. McKenzie, N., Coughlan, K. and Cresswell, H. (2002). Soil physical measurement and interpretation for land evaluation. CSIRO Publishing, Melbourne. Scotter, D.R., Clothier, B.E. and Harper, E.R. (1982). Measuring saturated hydraulic conductivity and sorptivity using twin rings. Aust. J. Soil Res. 20: 295-304.

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Infiltration Test Results

Infiltration at plot 2

4 test 1 test 2 3

2 Infiltration (cm) Infiltration

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

Infiltration at plot 22

test 1 3 test 2

2 Infiltration (cm) Infiltration

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

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Infiltration at plot 32

4 test 1 test 2 3

2 Infiltration (cm) Infiltration

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

Infiltration at plot 38

4 test 1 test 2 3

2 Infiltration (cm) Infiltration

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

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Infiltration at plot 47

4 test 1 test 2 3

Infiltration (cm) Infiltration 2

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

Infiltration at plot 62

2.5

1.5 test 1 test 2

1 Depth (cm) Depth

0.5

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

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Infiltration at plot 70

3.5

2.5

test 1 2 test 2

Depth (cm) Depth 1.5

0.5

0 0 5 10 15 20 25 30 35 40 45 50 55 60 Time (mins)

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