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SID 5 Research Project Final Report

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NoteIn line with the Freedom of Information Act 2000, Defra aims to place the results of its completed research projects in the public domain wherever possible. The SID 5 (Research Project Final Report) is designed to capture the information on the results and outputs of Defra-funded research in a format that is easily publishable through the Defra website. A SID 5 must be completed for all projects.

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Project identification

1. Defra Project code BD1444

2. Project title

Potential for enhancing biodiversity on intensive livestock farms

3. Contractororganisation(s)

Institute of Grassland and Environmental Research      British Trust for Ornithology     Centre for Agri-Environmental Research, University of Reading     

54. Total Defra project costs £ 1050588(agreed fixed price)

5. Project: start date................ 01 February 2002

end date................. 31 March 2007

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6. It is Defra’s intention to publish this form. Please confirm your agreement to do so...................................................................................YES NO (a) When preparing SID 5s contractors should bear in mind that Defra intends that they be made public. They

should be written in a clear and concise manner and represent a full account of the research project which someone not closely associated with the project can follow.Defra recognises that in a small minority of cases there may be information, such as intellectual property or commercially confidential data, used in or generated by the research project, which should not be disclosed. In these cases, such information should be detailed in a separate annex (not to be published) so that the SID 5 can be placed in the public domain. Where it is impossible to complete the Final Report without including references to any sensitive or confidential data, the information should be included and section (b) completed. NB: only in exceptional circumstances will Defra expect contractors to give a "No" answer.In all cases, reasons for withholding information must be fully in line with exemptions under the Environmental Information Regulations or the Freedom of Information Act 2000.

(b) If you have answered NO, please explain why the Final report should not be released into public domainN/A

Executive Summary7. The executive summary must not exceed 2 sides in total of A4 and should be understandable to the

intelligent non-scientist. It should cover the main objectives, methods and findings of the research, together with any other significant events and options for new work.Research to alleviate the decline of farmland fauna and flora has, to date, largely focussed on arable systems yet grasslands are an important habitat for many farmland species. Since the 1950s agricultural intensification has caused declines in many plant, insect and bird species associated with farmed landscapes. In grasslands the mechanisms responsible for the faunal declines remain poorly understood. The majority of agricultural grassland is now species poor and structurally uniform due to fertilisation, increased stocking levels and a switch from hay making to silage production. It appears axiomatic therefore that practices that create or enhance variability in vegetation structure and architecture within grassland habitats are likely to be beneficial to both birds and invertebrates. The overall aim of this project was to quantify the effects on botanical and faunal diversity of creating such heterogeneity, in intensively managed grassland farms, by applying simple management techniques to field margins. Specific objectives were: 1) Evaluate the effects of vegetation structure, height and species composition on invertebrate and bird populations 2) Determine how the experimental treatments differed in terms of numbers and species composition of plants, insects and birds. 3) Identify practical options for enhancing biodiversity on intensively managed livestock farms and 4) determine the cost-benefits of applying these different margin treatments. A randomised block design of nine different field margin treatments was established on two intensively managed grassland farms in Devon and two in Somerset. The farms were selected in areas with established populations of target farmland bird species e.g. Yellowhammer (Emberiza citrinella). The treatments were designed to provide a sequence of increasing structural and compositional complexity. They ranged from relatively simple practical options that farmers might adopt for a margin such as cessation of fertiliser application, raising mowing height, grazing leniently, or delaying cutting date. The most radical option applied to the existing grassland was to leave it uncut/ungrazed throughout the summer. Two sown treatments were included to provide greater variation in canopy cover, architecture and food resources for invertebrates and birds. One of these treatments involved a spring sown cereal undersown to a structurally diverse legume rich ley. A more complex mix of plant species designed to provide a wide range of seed and nectar resources was established in the second sown treatment. To ensure against climatic variation in establishment, both treatments were resown in each of three years giving three temporal replicates on each farm. Only the first temporal replicate of ley from the undersown cereal was maintained for the project’s duration.

Measurements carried out on each treatment replicate included: botanical surveys of plant species cover, canopy structure, counts of the seed and nectar resources. For the invertebrates: vortis sampling, sweep nets, pitfalls and timed transects were used to collect beetle, true-bug, planthopper, spider, grasshoppers, crane, St Marks fly, larvae (Lepidoptera and Symphyta), butterfly and bumblebee data. Timed counts of all bird species present on treatment plots, adjacent field boundary and adjoining field were made throughout each of the study years.

Ecological Response The dominant plant species on the existing grassland at the start of the experiment was ryegrass (Lolium perenne). During the course of the study the abundance of ryegrass declined particularly in the unfertilized, extensively managed treatments, being largely replaced by creeping bent (Agrostis

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stolonifera). The botanical diversity in the sown treatments was enhanced by the establishment of a wide range of unsown ruderal species, a notable example being sharp-leaved fluellen (Kickxia elatine). As intended the sown treatments provided a greater diversity and abundance of seed and nectar resources than the grass based treatments. No significant differences in plant species diversity were found during the course of this study between the treatments established on the existing grassland. Not surprisingly the sown treatments were significantly more diverse than the fertilized or the more frequently cut treatments, based on the original grassland. Dry matter yield (DMY) was significantly greater in the unfertilised grass treatment with one July cut compared with conventional management. However quality, both in terms of %N and digestible organic matter (%DOMD) was significantly greater in the conventionally managed treatment.

ConclusionTo date few studies have demonstrated biodiversity benefits from manipulating the structure and composition of grassland margins on intensively managed livestock farms. We show that sowing grassland margins with crop species to provide nectar, pollen and seed resources provided a rapid means of enhancing the numbers and diversity of birds and some insect taxa (particularly the pollinators) on intensive livestock farms. However, though sown options met their objectives in the short-term, they also presented longer-term management issues by allowing the ingress of pernicious weeds. We found that the establishment of the sown mixtures was variable both within and between the four farms, and this appeared to be associated with the soil conditions at the field margin. The soil was often compacted, had poor drainage and, where there was an adjacent ditch, commonly contained stones and other debris cleared from the ditch. These factors made preparation of a seed bed difficult and almost certainly contributed to patchy establishment. Areas of bare ground were readily colonised by pernicious weeds e.g. creeping thistle (Cirsium arvense). Any increase in pernicious weeds in field margin plots was perceived as a severe threat by the farmers/landowners to the adjacent crop. The uptake by farmers of management options that pose a weed threat is therefore likely to be limited.

The response of invertebrate and birds to the grassland treatments varied between species. The more extensive management treatments on the original grass, which received either one annual cut or no cut or grazing and no additional nutrients, supported the greatest invertebrate diversity in comparison with the conventionally managed control treatment, which was fertilized received two cuts and aftermath grazed. Although the result was much less consistent, the most extensive grass based treatment also allowed increase in some plant positive indicators of nature conservation value, for example, Devils’bit scabious (Succisa pratensis), and sneezewort (Achillea ptarmica).

Bird numbers on all farms were relatively low, reflecting both the impoverished bird communities in many grassland regions of the UK, the national declines of some these species and the small scale of the treatments for this taxa. Within the grass treatments (1 to 7) there was a general increase in use by small insectivores, e.g. Dunnock, with a decrease in management intensity of the sward, but the reverse was true for large insectivores, e.g. Blackbird. This almost certainly reflected a trade-off between prey abundance and accessibility, with small insectivores favouring the insect-rich taller swards and large insectivores favouring shorter swards where invertebrates were, generally, less abundant but more accessible. The sown swards (8 and 9) exhibited high use by granivorous finches and buntings in winter, reflecting the high seed resource these swards offer. They were also used by insectivores, at a higher frequency than grass swards, in winter and summer, despite the fact that these swards supported generally similar invertebrate abundances as the control grass treatment. Their high use probably reflects the fact that prey is more accessible in these patchy open swards compared to the taller denser grass swards present in treatments 5-7. The results show that grass swards managed relatively extensively, in relation to fertiliser input, cutting and grazing, will provide enhanced foraging resources for a range of bird species throughout the year. They could be provided as margins, part or whole fields and created by reducing fertiliser input as well as the cutting frequency and/or grazing intensity. However, the value of such grassland to most bird species is likely to be greatly enhanced if the relatively tall dense swards can be ‘opened up’ in a way that provides patches of short sward or bare ground. This will increase the accessibility of invertebrate prey to a range of insectivorous birds. Sown swards, particularly in the form of a wild bird cover crop but also as an undersown cereal, provide good foraging habitats for granivorous and insectivorous birds in winter, as do grass swards left to set seed in the first winter. Seed resources are rare in grassland in winter and even small areas of, for example wild bird cover crop, are likely to be used by relatively large numbers of birds throughout the winter.

This study has demonstrated that by using practical and low cost grassland extensification techniques biodiversity can be significantly enhanced on intensive livestock farms without having a major impact on other farm operations. Undisturbed or minimally disturbed field margins can fit in to typical livestock farming systems. Extensification practices that provide greatest biodiversity gain will, however, provide little if any forage of value to productive livestock. The scale of management practices required to enhance bird populations remains unknown, as does the relative cost effectiveness of these treatments at the whole field relative to the individual field margin scale.

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Options for New WorkFurther research is required to determine the optimal size and configuration of patches within grass swards such as those in undisturbed during the summer (treatment 7) to benefit a range of different taxa including insects and birds. The most cost-effective methods of creating such patches also need to be investigated. Introducing forbs into the grassland had both desirable and negative effects. More research is required on maximising the potential of introducing forbs into grassland either through slot-seeding techniques or more innovative methods. It is evident that some of the grassland margin treatments were more successful than others in enhancing biodiversity. Placing the most ‘beneficial’ treatment in a wider landscape could help to maximise local benefits.

Project Report to Defra8. As a guide this report should be no longer than 20 sides of A4. This report is to provide Defra with

details of the outputs of the research project for internal purposes; to meet the terms of the contract; and to allow Defra to publish details of the outputs to meet Environmental Information Regulation or Freedom of Information obligations. This short report to Defra does not preclude contractors from also seeking to publish a full, formal scientific report/paper in an appropriate scientific or other journal/publication. Indeed, Defra actively encourages such publications as part of the contract terms. The report to Defra should include: the scientific objectives as set out in the contract; the extent to which the objectives set out in the contract have been met; details of methods used and the results obtained, including statistical analysis (if appropriate); a discussion of the results and their reliability; the main implications of the findings; possible future work; and any action resulting from the research (e.g. IP, Knowledge Transfer).

ObjectivesThe overall aim of the project was to establish whether plant, insect and bird diversity could be enhanced within an intensively managed grassland landscape, by creating field margins which differed in terms of the vegetation composition, structure and height. Specific objectives were: 1) Evaluate the effects of vegetation structure, height and species composition on invertebrate and bird populations. 2) Determine how the experimental treatments differed both in terms of numbers and species composition of plants, insects and birds. 3) Identify practical options for enhancing biodiversity on intensively managed livestock farms. 4) Determine the cost-benefits of applying these different margin treatments.

All of these objectives were met.

MethodsTreatmentsFour intensively-managed grassland farms, all on clay soils, were selected in at two localities: Devon (North Wyke SX660984 and Heywoods SS642037) and Somerset (Bickenhall ST288191 and Southhill ST253182).

A randomised block design of nine different field margin treatments (Table 1) were established in April 2002 on each farm on permanent pastures (i.e. leys more than 5 years old) as this is the most common agricultural grassland habitat in the UK. Treatments were established in the field margins as bird species preferentially forage in and around hedgerows and invertebrates can readily colonise grasslands from them. The treatments measured 50m in length and 10m in width and had three replicates per farm.

The sown treatments 8 and 9 were designed to provide greater variation in canopy cover, architecture and food resources for the target invertebrate and bird community than could be achieved by manipulating the existing grass sward. To account for any climatic variation in establishment, both of these treatments were re-sown in each of three years giving three temporal replicates on each farm (table 2). Only the first temporal replicate of treatment 8 was maintained for the project’s duration. In order to reduce competition to the establishing kale seedlings the treatment 9 plots were divided in to two (5m x 50m) and the half closest to the hedge was sown with the quinoa and kale and the outer half to the cereal, linseed and legume mix.

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MeasurementsVegetation studiesBotanical compositionIn 2003 - 2006, a visual estimate of percentage cover of all vascular plant species, dead vegetation and bare ground was recorded in August on five 1m x 1m quadrats placed at set distances along a diagonal from the hedge to the field edge within each treatment plot. Each quadrat position was fixed throughout the experiment. Each plot was searched and any ‘additional species’ to those found within the fixed quadrats were recorded.

Table 1 Description of the field margin treatments implemented at four farms.  

Cutting height GrazedAug/Sep

Treatment N Fertiliser May July 1 Conventional silage management √ 5cm 5 cm √ 2 Unfertilized 5 cm 5 cm √ 3 Raised mowing height √ 10 cm 10 cm √ 4 No aftermath grazing √ 5 cm 5 cm 5 Single early cut 10 cm 6 Single late cut Hay cut in July 7 No summer disturbance Topped in Feb 8 Under-sown spring cereal Spring barley undersown with grass/legume mix†, cut July of

following year and subsequently managed as treatment 6

9 Sown complex mix Kale/quinoa/mixed cereal (barley, triticale, oats), linseed, legumes*. Topped in February.

†(cock’s-foot (Dactylis glomerata), brown bent (Agrostis capillaris), timothy (Phleum pratense), red fescue (Festuca rubra ssp Litoralis), crested dogstail (Cynosurus cristatus), sweet vernal (Anthoxanthum odoratum), ryegrass (Lolium perenne); red clover (Trifolium pratense), white clover (T. repens), bird’s-foot trefoil (Lotus corniculatus), common vetch (Vicia sativa) and black medick (Medicago lupulina ).* red clover (T. pratense), white clover (T. repens), bird’s-foot trefoil (L. corniculatus), vetch (V. sativa)

Table 2 Timings for sown field margin treatments (8 and 9).No Date Sown Date Cut/topped Date removed from

Experiment8a April 2003 July 2004, 2005, 2006 Sept 20079a April 2003 Feb 2005 Sept 20048b April 2004 July 2005 Sept 20059b April 2004 Feb 2006 Sept 20058c April 2005 July 2006 Sept 20069c April 2005 Sept 2007 Sept 2007

Vegetation Structure:A ‘drop disk’ method was used to assess leaf and stem density within the sward canopy. A wooden disk (30cm diameter, mass 200 g) was dropped from a height of 1m down a vertically held ruler and the resting height of the disc recorded. This provided a coarse estimate of vegetation structure. Twenty five regularly spaced drop disc measurements were made in April, June, July and September (to coincide with invertebrate sampling) along a diagonal from hedge to field boundary in each plot.

Fine grain vegetation structureA finer method was used in order to quantify in more detail the accessibility of the vegetation structure and composition to invertebrates. Ten point-quadrat pins, measuring 3mm in diameter, were placed at 10cm intervals along a 1m ruler. The ruler was thrown twice at random within each treatment and information gathered from a total of 20 pins per treatment replicate.

The pins were marked with height-category bands at 5cm intervals, and the number of ‘hits’ of each vegetation type within each band was counted. Vegetation type was classified as dead, legume, grass, forb, cereal or kale.

Each treatment 9 plot had two distinct halves, one with kale and quinoa and the other with the cereal, legume and linseed mix. Therefore 2 replicates i.e. a total of 20 pins were measured in each half in 2003 to 2004. From 2005 – 2006, because of the time taken to collect the data from a total of 36 treatment plots per farm (12 treatments (1-7, 8a, 8b, 8c, 9b, 9c) x 3 reps), the number of replicates of the pins was reduced on treatment 9 plots from 4 to 2: one in the kale and quinoa and the other in the cereal legume half.

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Seed and nectar resourcesIn 2003 - 2006, annual counts of the seed and nectar resources in each replicate were completed once in June (after the first silage cut) and again in September (after the grazing treatment), using circular quadrats 30cm in diameter. In 2003, 15 quadrats were thrown per treatment in June. From September 2003 through to the end of the experiment this number was reduced to 10 quadrats per treatment. In each instance the number of flowers/seed heads per species, rooted within the quadrat was recorded. Each species was then categorized into one of three functional groups: graminoids (grasses, rushes and sedges), forbs or legumes.

Agronomic measuresYield and quality (% nitrogen and % digestable organic matter (DOMD)) data for the harvested forage were collected from treatments 1-6 and treatment 8 (a, b and c) at the time of cutting. Two sub-samples of herbage were collected from a known length of swathe and the fresh weight of each sample recorded. Each sample was then dried in an oven at 850C before being ground for subsequent chemical analysis.

Invertebrate Sampling MethodsSeveral complementary methods were used to collect invertebrates in order to efficiently sample the full range of taxa from the vegetation, soil surface and below the ground:

(a) Vortis suctionA Vortis sampler (manufactured by Burkard®) was used to collect 5 samples from each plot. Each sample comprised fiveteen 10s sucks (total area 0.174m2) made by moving the Vortis vertically down onto the vegetation. Samples were evenly spaced out along the treatment replicate. Invertebrates were removed from debris by pooting and stored in 70% alcohol. Samples were collected in April, May, July and September.

(b) SweepingTwo 10m long transects, comprising 20 sweeps, were made on each side of a treatment replicate using a standard sweep net (Watkins and Doncaster). All Symphyta and Lepidoptera larvae were counted and weighed. Orthoptera were counted and identified where possible in situ. Samples were collected in April, May, July and September. An additional survey, in summer 2006 at Bickenhall, was used to assess the potential movement of Orthoptera from margin treatments into the field. A 50m long transect, parallel to the boundary, was made at 0, 5, 10 and 15m out from each treatment edge and at the centre of the field. From each sweep all Orthoptera were counted and identified.

(c) Pitfall trapsFive pitfalls were placed evenly along the centre of each treatment replicate and left open for two week period in March of each year. Pitfall traps comprised 60 mm diameter tubs (A W Gregory & Co Ltd) and filled with c. 100 ml 50% ethylene glycol and unscented detergent mix. Samples were frozen and retained for later identification in the lab.

(d) Pan traps for slugsTwo pan traps were evenly spaced along the centre line of each treatment replicate and collected after 2 days and slug numbers counted. Traps are 150 mm inverted flowerpot saucers baited with bran. Samples were collected in April, May, July and September.

(e) Timed transect walksA permanent transect route running along the centre line of treatment each 50m plot was walked (at 15-20m/min) once on each sampling date to count bumblebees and butterflies within 2.5m of the recorder. Walks were carried out between 10.00 and 17.00, when weather conformed to Butterfly Monitoring Scheme standards (i.e. temperature >13°C with at least 60% clear sky, or 17°C in any sky conditions, no rain, and wind speed <5 on the Beaufort scale). Transects were made in June, July, August and September.

(f) Below-ground macro-invertebratesBaseline data was collected in 2003 at one site using the electrical octet method. Subsequent calibration of this method allowed a detailed assessment of below-ground arthropods to be carried at another site in 2006 using hand-sorting of soil cores. In each grass treatment three cores (25 x 25cm area by 10cm deep) were extracted and all macro-invertebrates collected and stored in 70% alcohol.

Bird studiesUse by birds of PEBIL margins and adjacent areasTimed foraging watches were carried out to quantify the frequency with which different species used the experimental margins. These watches were carried out in three summers and three winters between November 2003 and October 2006, following pilot methodological work in year one (2003). Each treatment area (defined as the 50 m x 10 m experimental margin plus adjacent boundary hedge and field) was observed for two to four one-minute periods per month. Margins were observed from a distance of approximately 10 m from the near corner, recording the location of all birds seen in the boundary, margin and adjacent field. Positions of birds seen within

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the 10 m wide experimental margin were recorded in four distance bands from the boundary (0.0-2.5 m, 2.5–5.0 m, 5.0-7.5 m, and 7.5-10.0 m), and those in the adjacent field in a further three distance bands (10-20 m, 20–50 m and >50m from the boundary). Any birds flushed during the approach to the area were recorded separately and a single transect was walked through the centre of the experimental margin after the count to flush any previously undetected birds. Watches were carried out from one hour after dawn and no later than one hour before dusk, and poor conditions (wind and rain) were avoided. The order in which farms and treatments were visited was randomised to avoid bias due to time of day.

In an attempt to gather further information on foraging behaviour by Blackbirds and Dunnocks nesting adjacent to PEBIL margins, a trial radio-tracking study was carried out in May-June 2005. Birds were captured with the use of mist nets erected parallel to the field boundaries where they were nesting, and then fitted with tail-mounted transmitters that constituted approximately 2% of body mass. Dunnocks were fitted with 0.35 g. transmitters (length 13 mm, width 7 mm, height 4 mm), and Blackbirds with 2.4 g. transmitters (length 7 mm, width 13 mm, height 22 mm). Tags were attached to the base of the central pair of rectrices using a small amount of glue and nylon thread. The positions of birds were calculated by triangulation, using bearings taken simultaneously by two observers from a circuit of fixed points. Fieldwork was carried out during two discrete five-day periods within three weeks after tagging; one before the May silage cut and one after.

Territory mapping of study areasBreeding season territory mapping was carried out in three years (2003, 2004 and 2006) in order to assess the extent to which (a) the species using the margin reflected the species composition of the ‘wider’ bird community and (b) year to year changes in margin use could be explained by changes in local breeding abundance. Each area comprised fields that contained experimental margins and all the boundaries of those fields, as well as neighbouring fields and boundaries to within a distance of 100 m. Methods used were the same as those employed in the ‘reduced’ Common Bird Census (Marchant 1983) using four rather than eight to ten visits (e.g. Amar et al 2005). Visits took place between early April and mid June; one by 20 th April (Visit ‘A’), a second by 10th

May (‘B’), a third by 30th May (‘C’) and the fourth by 20th June (‘D’) each year

Nest site selectionDuring regular visits in April-June 2006, the locations of as many nests as possible in experimental margins and adjacent boundaries were identified. Search time was standardised, with equal time spent on each farm and each experimental margin within farms. Nests were located using a combination of “cold searching” and using obvious signs of breeding activity. Height of the nest (to nearest 0.1 m), plant species present in the immediate vicinity and an assessment of exposure (well hidden, part hidden, or exposed) were recorded for all nests found.

Boundary structure characteristicsMargin use by birds is heavily influenced by the nature and extent of adjacent boundary and to control for this potentially confounding effect, boundary characteristics were measured in August 2003 and 2006. These included height, width (recorded to the nearest 0.1m at 10m intervals along the hedge), proximity of mature trees and presence of herbaceous vegetation at the boundary-margin interface.

Statistical AnalysisThe data was analysed as a two-way ANOVA, where the explanatory factors were i) treatment and ii) farm nested within site. In order to take into consideration the different starting point of the sown treatments, in comparison with the seven grassland treatments, each year was analysed separately. The experiment can be considered as two separate systems: manipulation of grassland field margins (T1-7) and the creation of sown margins (T8 and T9). The grass system (i.e. T 1-7) is not disturbed in anyway and so the whole system including the soil remains intact for the project’s duration. Creating the sown treatments disrupts the grass system, exposing bare soil. Subsequently for each year, there was an analysis containing all seven grass treatments wand another analysis comparing differences between sown treatments created in the same year. Significant differences in the response variable e.g. species number between a sown treatment e.g. T8a and the control T1 were compared by means of a t-test.

Vegetation and invertebrate data Statistical Analysis was performed using the freeware R package version 2.4.1. All data relating to the vegetation was transformed using the natural log except where stated. Where transformation failed to normalise the residuals of the data a non-parameteric Kruskal-Wallis test was performed on the un-transformed data.

Changes in botanical composition were analysed using RDA analysis in CANOCO 4.5 (Braak & Smilauer, 2002). In all cases the data was log transformed prior to analysis. Only those species which had on average more than 1% cover from all five quadrats were included in the analysis.

Bird dataForaging counts: the data were analysed by season (winter: October to March; summer April to September) and logistic regression was used to determine treatment effects. We used an events/trials syntax where the events were the number of one minute counts in which the species was present in the margin or adjacent boundary and the trials were the total number of counts under taken each season. The dependent variables were treatment,

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county, farm and farm*county and county*treatment interactions. Nest distribution: this could be affected by the characteristics of the hedge (structure and vegetation composition) or the adjacent margin. We therefore first tested for differences in hedge characteristics in relation to treatment, farm and an interaction term to determine whether there was systematic bias between treatments and farms. We then tested whether nest distribution was related to farm and (a) hedge characteristics and (b) the adjacent treatment type. A generalised linear model (with appropriate link functions and error structure) was used to test for these differences. Normal errors & an identify link function were used to test for differences in hedge structure (height and width) between treatments. To identify whether nests were associated with particular plant species, we divided up each 50m plot into five 10m sections and recorded whether hawthorn, blackthorn, bramble, dog rose and nettle were present, these being identified as relevant to songbird nest site selection within hedgerows. To determine whether there was bias between treatments we used an events/trials syntax such that the number of events were the number of 10m sections a species was recorded in and the trials were always five, being the number of sections each plot was divided into. A logit link function and binomial errors were specified. The relationship between the number of nests vs treatment was tested for using a Poisson error structure and a log link function. Adjacent margin type was categorised as either “sown” or a “grass ley”. A sown margin was defined as one not being managed as a typical grass ley in April-June 2006 – and hence referred to all replicates of treatments 9b, 9c and 8c. Though 8b and 9b were both sown in 2004, 8b was undersown with grass and consequently managed as a grass ley by spring 2006. Consequently, for the purposes of this analysis each farm contained three replicates of 9 ‘ley margins’ (treatments 1-7, 8a-b) and 3 ‘sown margins’ (8c, 9b-c). (Treatment 9a was no longer in situ in 2006). This equates to 1350 m of grass ley margin and 450 m of sown margin per farm, or 5400 m of grass ley margin and 1800 m of sown margin across all four farms.

ResultsVegetation Results for vegetation response within the field margin treatments presented below are a condensed version of those presented within Appendix 1. Both county (Somerset and Devon) and farm had significant effects, at least within individual years, for most of the vegetation data considered. Regional differences in the species pool and historical differences in farm management probably contributed to the observed high variation in the botanical data between sites. The results presented below focus on overall treatment effects.

Species Number: Across all years there was no significant difference in species number between the seven treatments based on the original grassland. As expected in the year of sowing the sown treatments, 8 and 9, there were significantly higher number of plant species in comparison with the control treatment 1. In their second year no significant difference between sown treatments and the control was found (Table A1, Figure A1).

Botanical Composition Grass treatments: Botanical composition at the beginning of the experiment (2003) was compared with that at the end (2006). In 2003 county (Devon vs Somerset) explained 6.4% of the total variation whilst farms within counties explained a further 15.7% of the variation. Including treatment as well in the analysis resulted in a total 32.9% of the total variation being explained. Using Monte Carlo simulation tests revealed significant differences between treatments in species composition (test on both canonical axes, F-ratio = 1.804, p<0.05). Those treatments which received fertiliser (treatments 1-4) had a very similar species composition, dominated mainly by Lolium perenne, Taraxacum officinale and Rumex acetosa in 2003 (Fig. 1a). These treatments also had a higher percentage cover of litter and bare ground in comparison with treatments 5-7. Treatment 6 differed the most between the two regions, with Devon being the treatment least similar to any of the others, due to its domination by Festuca rubra and Holcus lanatus. It is probable that soil wetness of particular plots was a factor making treatment 7 in Somerset and treatment 5 in Devon appear to be botanically close.

In 2006 a total of 40.6% of the total species variation could be explained (county 7.8%, farm 15% and treatment 17.8%). Significant differences were only found between treatments (test on both canonical axes = 0.23, F-ratio=2.44, p<0.01).

Clear separation in the species composition between treatments which received fertiliser (treatments 1-4) and those which did not (treatment 5 & 7) was observed in 2006 (Fig. 1b). The unfertilized treatment 6, however, had a similar species composition to some of the fertilised plots such as, for example, treatment 3 in both regions and treatment 4 in Devon, all of which had a high percentage cover of Holcus lanatus and bare ground. Generally the fertilised treatment plots in 2006 had a species composition which was very similar to that which they had in 2003, namely high in Lolium perenne and Taraxacum officinalis with a greater percentage cover of bare ground and litter. The unfertilised treatments 5 and 7 were characterised by having a higher percentage cover of Agrostis stolonifers, Ranunculus repens, Arrhenatherum elatius and Cirsium arvensis.

For the sown treatments in 2003, 8a and 9a, the percentage of variation explained can be broken down into farm 10.5 %, site 12.5% and treatment 29%. The differences between treatments largely reflect the differences in the sown mixtures (Appendix 1, Fig. A2a). However, some of the differences between the two sown treatments reflect differences in the ruderal species associated with them. For example, there was more Cirsium arvense in treatments 9a in Devon and 8a in Somerset, whilst treatment 8a in Devon had a greater percentage cover of Persicaria maculosa, and Geranium dissectum. In 2004 county only explained 9.2% of the variation, farm accounts for a further 11.3% with treatment explaining 28.1%. Significant differences remain between the two sown treatments (trace = 0.249, Fratio=8.2, p<0.05). Treatment 8a had become dominated by the sown grasses Dactylis glomeratus, Anthoxanthum

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odoratum and Festuca rubra and the sown legumes Medicago lupulina and Lotus corniculatus and bare ground (Fig. A2b) . Treatment 9a was characterised by fewer species in its second year. Pernicious weeds such as Cirsium vulgaris and Urtica dioica were present as well as the ‘sown species’ Kale. Similar results were found for treatments 8b, 9b, 8c and 9c. The ordination diagrams are presented in the appendix (Figs. A3 and A4).

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Figure 1a Species composition of the 7 grassland treatments within the two different counties in 2003. The species fit range is 5-100%. b) the same 7 grassland treatments at the end of the experiment in 2006. Species fit range is 7 -100%.

By 2006 farm and county explained 39.1% of the total variation in species composition of treatment 8a. The species composition of this treatment was not found to differ significantly either between counties (test on both canonical axes = 0.2, Fratio=24.4 p=1) or farms (test on both canonical axes =0.24, F-ratio=1.59, p>0.05)

Seed and nectar resources (Tables A2-A4): Very few if any flower or seed heads were recorded in the plots that were cut twice and this presented a large numbers of zeros in the data set. Flower and seed head data analyses were restricted to those treatments which had a specified number of seed heads per m 2 e.g. >5 or in some cases >0, in order to normalise the residuals.

Graminoids: Grass treatments As expected in all years, there were significant differences in the number of graminoid seed heads between the seven treatments based on the original grassland. Treatments which only received one cut between May and July (treatments 5 and 6) or no cut (treatment 7) had significantly more graminoid seed resources (Fig 2a).

Graminoids: Sown treatments; graminoid seed head numbers were significantly higher in the grass dominated treatment 8 plots in comparison with treatment 9. In all years treatment 8a had significantly more seed heads than the control treatment 1 (Fig. 2b).

Legumes: Grass treatments. Numbers of seed heads of legumes were very low in the grass treatments for all years. No significant differences in legume resources were found between treatments 1-7 (Fig 2c).

Legumes: Sown treatments. Contrastingly legume resources were much higher in the sown treatments (Fig 2d), though this is not surprising as they were specifically sown in both treatments 8 and 9. Treatment 8a had significantly more legumes than the control (2003-2005) and 9a (2003 and 2004), 8b (2004). In 2006 treatment 9c had significantly more legume resources in comparison with 8c, though neither 8c nor 9c had significantly more legume resources than the control in 2006.

Forbs: Grass treatments. Forb numbers on the grass treatments were low. In 2003 a Kruskal-Wallis analysis showed that there were significantly more forbs in treatment 6 compared with the other treatments. 2006 was the only other year where significant difference in forb seed resources was found between treatments, with treatments 4,6 and 7 having significantly more forbs than the other treatments.

Forbs: Sown treatments. As expected, forb numbers were higher on treatment 9 compared with treatment 8 in the year the treatments were sown for both 9a and 9b. Though 9a continued to have significantly higher forb resource than 8a in its second year, no significant difference was found between 9b and 8b in 2005. No significant difference in forb resource was found between 8c or 9c for either year. Not surprisingly all the temporal sown replicates of both treatments 8 and 9 had significantly more forb seed heads than the control (treatment 1).

Agronomic measures (Tables A5-A7): DMY/kg/ha: Grass treatments There were some significant differences between treatments within counties and between counties for some of the years (table A5). In general there were significant differences in silage yield between treatments: Treatments 5 and 2 had a tendency to be lower in yield compared to the control whilst treatment 6 had a significantly higher yield (fig 3a).

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Figure 2: Nectar and seed resources on both grass and sown treatments a) graminoids in the grass treatments, b) graminoids in the sown treatments, c) legumes in grass treatments, d) legumes in sown treatments , e) forbs in grass treatments, f) forbs in sown treatments NB note the different scales between the grass and the sown treatments for forb resources. Kruskal-Wallis test was used for analysis of the forb data (treatments 1-7 and 8a and 9a 2003) and legume seed resources (treatments 8a and 9a 2003) so no mean is shown.. Please refer to Appendix I tables A3 and A4.

Dry matter Yield (DMY/kg/ha): Sown treatments None of the temporal Treatment 8 replicates had significantly higher silage yield compared to the control treatment (Fig 3b).

%N:Grass treatments As for DMY there instances of significant differences in %N for the same treatment but on different farms within a county. Generally %N was significantly lower in the unfertilised treatments, especially treatment 6, compared with treatments 1-4 (Fig 3c).

%N Sown treatments The %N of the cut herbage was significantly lower in all of the treatment 8 temporal replicates in all years (Fig 3d).

%DOMD:Grass treatments Across all three years, treatment 6 had significantly lower %DOMD in comparison with the control treatment 1 (Fig 3e)

%DOMD:Sown treatments. For all years the temporal replicates of treatment 8 had significantly lower %DOMD in comparasion with the herbage cut from the control.

InvertebratesResults for invertebrate response within the field margin treatments are a condensed version of those presented within Appendix 2. Both county (Somerset and Devon) and farm had significant effects, at least within individual years, for all of the invertebrate taxa considered. However, such between site variations was expected given both regional differences in the species pool and historical differences in farm management. The focus of this results section therefore is on between treatment differences in margin management.

SID 5 (Rev. 3/06) Page 11 of 23

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Figure 3. Agronomic measures on both the grass and sown treatments a) Dry matter yield (DMY kg/ha) in treatments 1-6 b) DMY kg/ha in temporal replicates of treatment 8 c) %N in treatments 1-6 d) %N in temporal replicates of treatment 8 e) % DOMD in treatments 1-6 e) %DOMD in temporal replicates of treatment 8

Beetles (suction sampling)(Fig 4) Margin treatment had significant effects on beetle abundance (all years) and species richness (2005 and 2006 only) for the grass-based plots (treatments 1-7). This pattern was characterised by high abundances in the intensively managed treatments, i.e. those receiving multiple silage cuts each year in combination with NPK fertiliser (treatments 1, 3 and 4). However, by 2005 - 2006 there were apparent successional shifts within the grass-based margins that were characterised by higher abundances of beetles within the extensively managed grass swards, particularly treatments 5, 6 and 7. A similar pattern was also found in overall beetle species richness, with the extensively managed grass manipulation treatments (5-7) supporting significantly higher species richness by 2005 and 2006. Generally, the sown treatments (8 and 9) were not found to support beetle abundances or species richness that differed significantly from the control treatment of the grass-based plots (treatment 1). Comparisons between the sown treatments 8 and 9 within individuals years showed that for both beetle abundance and species richness significant differences between these treatments were infrequent.

Ground beetles (pitfall trapping): Ground beetles collected using pitfall traps in March showed no between treatment differences for the grass-based plots in either activity abundance or species richness.

Bumblebees: Total numbers of bumblebees were highly variable between years and between farms. Within the grass-based plots, there were significant treatment differences for abundance in 2004 and 2006, where treatment 6 supported the greatest number of bumblebees. Treatments 8 and 9 always had more bumblebees and greater species richness than the control treatment for the grass-based margins (treatment 1); however, the abundance trend was only statistically significant for treatments 8 and 9 in 2004, 9b in 2005, and 9c in 2006.

SID 5 (Rev. 3/06) Page 12 of 23

Within most years, treatments 8 and 9 performed equally well in terms of bumblebee abundance and diversity, though there were some exceptions.

Butterflies: Total numbers of butterflies fluctuated between years and farms, but were much less variable than bumblebees. There was a significant, and generally consistent, treatment effect on butterfly abundance for the grass-based treatments in all years, where treatments 6 and 7 supported greater numbers of butterflies. Butterfly species richness was also significantly affected in years 2004 to 2006 and showed a broadly similar pattern to butterfly abundance. In all years, the 8 and 9 treatments consistently had greater butterfly numbers, and almost always more butterfly species than treatment 1 (Fig. 5). With the exception of 2004 treatments 8 and 9 were very similar in abundance and species richness.

True bugs: For the grass-based plots (treatments 1-7), treatment had a significant effect on true bug abundance; the general pattern was for treatments 1-4 to support lower numbers than treatments 5-7. Species richness was also significantly affected by grass treatments, with the greatest diversity in taller grass swards in treatments 5-7. Abundance of true bugs for both sown treatments 8 and 9 was generally lower than that of the control treatment, although this pattern was not seen for true bug species richness. Treatments 8 and 9 on the whole did not differ in either true bug abundance or species richness.

Plant hoppers: There were consistent between treatment differences for the grass-based margins in most years for both abundance and species richness. Both treatment 1 (the control) and treatment 2 tended to support lower plant hopper abundances; however, species richness was generally greatest across all years for the extensively managed treatments 5-7. Both plant hopper abundance and species richness in the sown treatments was for the most part lower than that of the control treatment of the grass-based margins. Treatment 8 of the sown margins often supported higher abundances and species richness than treatment 9, although not in all years.

Spiders: For the grass-based plots the extensively managed treatments 5 and 7, as well as treatment 4 supported the greatest abundances of spiders, although the strength of this pattern varied between years. There was no evidence for differences in the abundances of spiders in the sown treatments 8 and 9 when compared to the control (treatment 1). When considered on their own, significant differences in spider abundance between the sown treatments 8 and 9 were largely absent. Orthoptera: For the grass-based margins there were no between treatment differences in Orthoptera abundance for any of the years, however there were large between site differences. Treatments 8 and 9 showed no consistent between treatment differences and neither was significantly different from the control treatment of the grass-based margins. There was no evidence that margin type had a significant effect on the distance Orthoptera dispersed into the field.

Flies: Significant treatment effects in 2003, 2004 and 2006 for the grass-based margins indicated that higher abundances of crane and St Marks flies were associated with the extensively managed treatments 5 -7. Although there was some evidence that relative to the control treatment the sown treatment 8 supported higher abundances of these flies in some years, this effect was not consistent and was entirely absent for treatment 9. Treatments 8 and 9 also showed no significant between treatment differences.

Larvae: Larval (Symphyta and Lepidoptera) abundance and mass responded significantly to the effects of treatment for the grass-based margins in 2003 and 2006, where both parameters were consistently lower in treatment 2. Larval abundance and mass in the sown treatments (8 and 9) did not differ from each other or from the control (treatment 1).

SID 5 (Rev. 3/06) Page 13 of 23

10.0012.0014.0016.0018.0020.0022.0024.0026.0028.00

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Fig 4. Response of beetle species richness from the Vortis suction samples to treatments 1 – 7 representing the management manipulations of the existing improved grassland sward.

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Slugs: Significant between treatment differences for the grass-based margins were only found in 2003, where there was a weak trend of higher abundances in treatments 2 and 7. Slug abundance of the sown treatments 8 and 9 did not on the whole differ significantly from each other or from the control treatment 1.

Below-ground invertebrates: For all below-ground taxa of invertebrates there were no significant differences in numbers between the grass-based treatments 1 to 7. For those groups for which data were available (Lumbricidae, Isopoda and Chilopoda) there was no difference in species richness, except for the Isopoda which were generally more diverse in the extensive treatments 5-7 (Appendix 3).

BirdsTerritory Analysis: The breeding species present within the four study areas (Appendix 4, Table A32) were typical of breeding bird communities of pastoral areas in lowland Britain. Eleven species were recorded breeding in all years in the fields and boundaries adjacent to experimental margins to within 100m (Fig.6a). Changes in territory density between the start and finish of the project (2003-2006) were generally small with six species increasing and five decreasing. The majority of species exhibited trends similar to those derived from the BBS data for south-west England for the period 1994-2005 (Fig 6b). Great Tit and Blackbird were anomalies, showing small declines in our study areas but increasing markedly within the south-west region. Reasons for these two differences are unclear. However, given our sample size of just four study areas, some apparent differences with respect to individual species population trends are probably to be expected.a) b)

Fig 6(a) Mean number (+SE) of territories per km of boundary in 2003 and 2006 of the 11 species recorded as breeding in all study areas throughout project; (b) Comparison of population trends of the 11 species recorded as breeding in all study areas throughout project with trends derived from regional BBS (95% CL)

Foraging counts: The generally low number of birds using the plots and adjacent boundary meant it was only possible to include county, farm and treatment in the final analyses. The small insectivores (Dunnock, Wren and Robin) were the most frequently recorded group in the grass treatments only and there was a significant treatment effect within these in summer 2004 and 2006 and in the second and third winters. (Table A34). No single treatment consistently supported the highest use by these species in summer but there was a marked linear increase along the management intensity gradient from treatment 1 to 7 (figure 9). In 2006, for example, small insectivores were recorded on c10% of visits in treatment 1 compared to c20% in treatment 7. The effect was less marked in winter but there was still an increase in use from treatments 2 to 7. Overall, small insectivores were most likely to be found in the sown treatments (10/11 comparisons between treatment 1 and sown treatments were significant). Large insectivores (thrushes and starling) were rarely encountered on the grass plots in summer (generally <5% of counts) and there were no clear treatment effects (Table A35). In winter, this group appeared to prefer the more intensively managed treatments, especially treatment 3. In summer, there was an approximate doubling in the frequency with which finches and buntings were recorded from treatment 1 to 7 and significant treatment effects in both years (Table A35). With respect to winter, there was a strong treatment effect in the first winter largely attributable to the very high use of treatment 7, but this disappeared in the subsequent winters. Notably, in the first winter, this treatment supported several flocks (5-20 individuals) of Yellowhammers feeding on the seeds of the rye grass. Of the sown treatments, treatment 9 (kale and quinoa) was used with the greatest frequency in its first and second winter. Treatment 8a (undersown cereal) was used to a lesser extent than kale/quinoa by granivorous birds. Its use diminished over the three winters and by the third year, when it had reverted to a grass ley, it was not significantly different from treatment 1.

In agreement with the standardised foraging counts, a radio tracking study at Heywoods Farm in spring 2005 indicated relatively low use of the experimental field margins by both Dunnocks and Blackbirds during the breeding period (a review was included in 2005 Annual Report). In summary, three individual Dunnocks preferentially foraged within bases of hedgerows or in adjacent scrub/garden habitats, and five Blackbirds made regular use of a manure heap sited on the edge of the study area. However, despite limited data, it was apparent that the Blackbirds made greater use of field margins for foraging after the grass had been cut for silage, indicated in Figs 7 and 8 below.

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Nest site selection: Only a single nest site (that of a Whitethroat in a 9c margin) was located within an experimental margin but a total of 72 nests of 13 species were located in adjacent hedgerows. These were, in order of abundance, Chaffinch (19), Blackbird (17), Dunnock (11), Goldfinch (7), Robin (5), Song thrush (4), Wren (3), Whitethroat (1), Chiffchaff (1), Willow Warbler (1), Greenfinch (1), Linnet (1) and Yellowhammer (1). As far as was possible to determine from territory mapping and foraging observations, all nests related to different individual territories and therefore do not include any repeat nesting attempts by the same birds. There were differences in the mean height and width of hedges between sites (χ2

3=65.63***) but not between treatment type (χ2

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between sown and ley treatments, with each species more likely to occur on ley treatments. There were sufficient data to test for the effects of treatment on nest site selection in three species (Chaffinch, Dunnock and Blackbird), although the small sample size meant that it was not possible to test for a site by treatment type interaction. No significant relationships were found between the number of nests of each species and (a) hedge height or width or (b) the proportion of sections of each treatment type occupied by Blackthorn, Dog rose, Hawthorn or Nettle (tests not shown). There was no site (farm) effect for any of the three species but a strong treatment effect was found for Dunnock (χ2

1=7.29**) with more nests than expected being found adjacent to sown treatments (Fig. 10), indicating that the adjacent margin type influenced nest site selection. Small insectivorous species such as Dunnocks have relatively small territories and will favour foraging close to their nest site. There was no significant increase in the density of Dunnock territories between 2003 and 2006 and it is likely that this was redistribution, rather than a genuine increase in Dunnock density related to altered margin management.

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Chaffinch (19) Blackbird (17) Dunnock (11)0.0

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DiscussionVegetation Plant species number either per m2 or per plot on the treatments based on the original grassland showed no effects of either cessation of fertilizer inputs or reduction in disturbance during the course of this study. This presumably indicated that factors such as residual fertility and recruitment niche space severely impeded change in botanical diversity of such vegetation. The sown treatments 8 and 9 were designed to overcome this spatial structural inertia and significantly higher plant species diversity were achieved in both treatments compared with the control treatment on the original sward. An additional bonus on the sown treatments was the recruitment of unsown ruderal species some of which are declining arable weed species.

Botanical composition was however, more responsive to treatment effects than species number, per se. Within the first growing season the composition of the unfertilized single cut plots diverged from those that continued to receive fertilizer, with one apparent anomaly, the hay cutting treatment, which remained similar in composition to the fertilized treatments as in 2003. By the end of the experiment, in 2006, the unfertilized undisturbed and May cut only treatments were distinctly different in botanical composition from the other treatments based on the original grassland. For example, a major component of the unfertilized plots that were either cut once, in May, or undisturbed throughout the summer was Agrostis capillaris, which in accordance with its Ellenberg N index (Ellenberg, 1988; Hill et al., 1999) is a less nutrient demanding species than L. perenne. The grass Arrhenatherum elatius, which is intolerant of frequent disturbance such as heavy grazing (Hubbard, 1972) was advantaged by the more extensive management treatments. The single cutting regime also appeared to favour the ingress of Cirsium arvensis from the hedge. Treatment 6, which was unfertilized and cut once in July, appeared to be anomalous with its species composition being similar to that of the fertilized treatments 3 and 4. It appears that the late cutting of treatment 6 allowed more competitive species to maintain an advantage over less nutrient demanding species during the early summer, particularly where the soils were relatively fertile. Where treatments involved two cuts dead plant material formed a greater proportion of the ground cover than in the more extensive treatments (where much of the dead leaf, stem and any flower/seed heads were removed by the harvest, except for the undisturbed treatment 7).

The sown treatments were successful in significantly enhancing the number of species present and thus spatial heterogeneity in field margins plots. However, the temporal replicates of the sown treatments did highlight problems in establishing these treatments. In 2003 the three replicates of treatments 8 and 9 established relatively well on all four farms, but in 2004 establishment was much more variable; more than 50% of the sown species established for only one replicate of treatment 9 in Somerset. In 2005 one replicate of treatment 8 failed to establish in Somerset. This poor establishment reflects the poor soil structure of the margins in grassland farms. These are the areas where farm machinery turns, where there is machinery trafficking when hedge trimming is carried out and is where debris from ditch clearances are placed. All four farms had clay soils and this coupled with poor soil structure, compaction and drainage problems made it extremely difficult to prepare a fine tilth for the seed-bed in time for optimum sowing conditions. Poor seed establishment created disturbed bare ground, which allowed ingress of opportunistic ruderal species such as Cirsium arvense and C. vulgaris.

The project showed that a greater number of seed resources could be provided from the original grassland simply by either cutting once in May and then leaving the grassland undisturbed (treatment 5) or by leaving the sward undisturbed throughout the summer (treatment 7). Cutting in July provided no seed resources in the autumn. Although treatments 5-7 provided seed resources for invertebrates and birds, these were predominantly from grasses, and this is likely to be the case for most de-intensified grassland. A much greater variety of seed and nectar resources were provided by the sown treatments, notably treatment 9, with a mixture of forbs (quinoa, linseed), graminoids (triticale, oats, barley) and legumes (white and red clover, birdsfoot trefoil and vetch). Treatment 8 provided a more limited range of pollen and nectar resources, with a mixture of grasses

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(Agrostis capillaris, Phleum pratense, Anthoxanthum odoratum, Lolium perenne, barley), and legumes (red and white clover, black medick, birdsfoot trefoil and vetch). However, differences in the variety of pollen and nectar resources were only found consistently for the first temporal replicates of treatments 8 and 9, which reflected the poor establishment of these subsequent sown treatments.

The potential agronomic value of the treatments that provided some agronomic return, i.e. were cut during the growing season, was assessed in terms of dry matter yield produced and the quality of the cut herbage. Cessation of fertilizer input to the field margin, but with continuation of conventional silage management, i.e. cut in May and in July, reduced dry matter yield by c. 2 t/ha and reduced N content by approximately 0.5 of a percentage unit compared with the fertilized control. However, simply stopping fertilizer inputs to the field margin had no effect on the digestibility of the forage produced. Raising the mowing height from 5 cm to 10cm on what were otherwise intensively managed grass margins reduced yield by c.1.2 t /ha, but had no effect on N content or the digestibility of the herbage compared with the conventionally managed control. These data, therefore, provide a measure of the agronomic penalty, which amounted to 30-40 % yield loss that would result from implementing what are relatively simple practical measures on the margins of intensively managed grass fields. The cut forage from the margin could be readily included in the forage from the rest of the field without compromising the overall quality of the silage; therefore the forage from the margin is readily usable by livestock farmers. However, the yield penalties need to be balanced against what were relatively modest biodiversity gains. The treatments that provided greatest biodiversity gains were those that involved a single cut or no cut during the summer. The unfertilized treatment involving a single cut in May reduced the total dry matter yield by c.2 t /ha, i.e. approximately a 40 % yield reduction compared with the control. There was also a reduced N content of the forage, but no effect on digestibility. Again the cut herbage from this treatment could readily be incorporated into the silage from the whole field. The depression in N (crude protein) content would be diluted within the bulk of the forage from the rest of the field and would probably have no significant overall effect on feeding value of the silage. Unfertilized plots that were cut once in July (treatment 6) produced c. 1.7 t /ha, c. 35 % more dry matter than the fertilised twice cut control. However, the nitrogen content and digestibility of the herbage were considerably reduced, (by c. 50 % and 10%, respectively) compared with the control. Nevertheless such forage would be adequate as part of the feed ration of dry cows or forward store cattle during the winter months. A similar argument could be made for the forage produced from the sown treatment 8 plots, whose yield and quality were similar to treatment 6.

InvertebratesThe discussion for the invertebrates is split into two parts considering the grass-based manipulation of existing sward (treatments 1-7) and the sown margins (treatments 8 and 9). For a number of taxa, specifically slugs and Orthoptera, treatments effects both of the grass-based and sown margin treatments were unimportant and they have therefore not been considered here.

Grass-based margins (treatments 1-7): The form of management used in the creation of the grass-based margins was for most taxa important in influencing both their abundance and species richness. The notable exception to this was the bumblebees, which were generally unresponsive to management treatments 1-7. However, large inter-annual variation is typical of bee communities, reflecting their high mobility and heterogeneous local spatial distributions (Williams et al., 2001), this may have impacted on the findings for the grass-based treatments.

For the beetles, butterflies, true bugs, plant hoppers and spiders (abundance only) management of the grass-based treatments had important impacts on both abundance and species richness. Although there was often between year variation in these trends, there was on the whole a consistent relationship between reduction in the intensity with which these grass-based margins were managed (in terms of NPK fertiliser application, grazing and frequency of silage cuts) and increasing abundance and species richness for these taxa. This trend of increased abundance and species richness was most apparent for treatments 5-7. There was also evidence in some taxa of a temporal increase in the quality of the margins from 2003 to 2006, e.g. the beetles. For this group the value of margins was not realised immediately, and instead required between 2-3 years before the benefits were fully realised, particularly in terms of the number of species supported in the extensively managed margins.

The general trend of increasing field margin value for the invertebrates, where extensive management regimes were applied (treatments 5-7), was likely to be the result of a combination of factors relating to individual functional requirements of the invertebrate species. Within the grass-based margins sward architectural complexity was likely to have had an important impact on both the structure and diversity of the invertebrate assemblages (Gibson et al., 1992; Woodcock et al., 2007; Woodcock et al., In press). For the extensively managed treatments 5-7, reduced frequency of cutting, as well as the absence of aftermath grazing, resulted in swards characterised by greater overall height and increased density of plant structures (foliage, stems, inflorescences and seed heads). The role that sward architecture plays in structuring invertebrate assemblages varies between the different taxa in this study. For example, in the case of phytophagous invertebrates (e.g. some beetles and true bugs, and all plant hoppers) the presence of host plants alone is not necessarily sufficient for their successful utilisation of the margins. Instead key plant architectural structures will be required by some species, e.g. seed heads for some Apionidae weevils. In contrast, for many phytophagous invertebrates that are successful within intensively managed grasslands their larval / juvenile stages are often associated with plant structures that remain largely undamaged by management practices such as cutting or grazing, e.g. root feeding

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weevils of the genus Sitona spp. and the ground-dwelling nymphs of Aphrodes spp plant hoppers. For phytophagous taxa the importance of sward architectural complexity will also interact with the diversity of the floral community. However, the establishment of new floral species, in particular forbs, within the grass-based margins was often slow. This relative lack of forb diversity, even in the extensively managed treatments 6 and 7, may explain the unexpectedly low species richness of many taxa, e.g. the butterflies, which were numerically dominated by the meadow brown in the grass-based margins. Sward architecture may also have limited the numbers of Symphyta and Lepidopteran larvae in treatment 2, as a simple product of a reduction in the availability of foliage on which to feed. This is consistent with the multiple silage cuts made in treatment 2, but in contrast to the other intensively managed treatments, the lack of NPK fertiliser meant that re-growth of the sward was comparatively slow. In the case of obligate predatory invertebrates (e.g. spiders and some beetles) most will have no direct links with specific plant species and will instead be dependent on general characteristics of sward architecture. For many spiders, architecturally complex swards provide web building structures, although the development of tussock grasses within the extensively managed treatments will also provide important refuges for many other predatory taxa. It should be noted that in the case of some plant species their may exist what could be considered as species specific associations shown by obligate predatory invertebrates, however, this normally occurs for plants that make an important or defining contribution to the architectural structure of the sward, e.g. Deschampsia or Dactylis spp (Poaceae). The Isopoda detritivore communities associated with the extensively managed grass-based treatments were also likely to have benefited from the increased sward architectural complexity of these treatments in terms of structural refuges and the greater accumulation of detritus/dead plant material. However, while significant responses to the grass based treatments were found for the Isopoda, the absence of responses for other bellow ground taxa is thought to reflect the limited sampling based on a single farm and sample date used for these groups.

Sown margins (treatments 8-9): For bumblebees, the more interventionist treatments of 8 and 9 consistently supported higher abundances and greater species richness than the control of the grass-based plots (treatment 1). The sown plots provided a much more diverse and reward-rich flower community than any of the grass treatments, which were characterised by few floral resources. In almost all cases the only floral resource found in any abundance within the grass-based treatments (1-7) was white clover. There is an indication that the benefits of the sown components for bumblebees are greatest in the second year after sowing, at least for treatments 8 and 9 in 2004. Longer-term benefits are unlikely, given that floral succession results in a loss of the sown species component which provides the best forage plants for bumblebees.

For most taxa the sown treatments were often characterised by not differing significantly for the control of the grass based margins (treatment 1). An exception was plant hoppers, which were consistently less abundant and species rich in the sown margins. This is likely to reflect the preference of most plant hoppers for grasses as their host plants, which were dominant in treatments 1-7, whereas the sown plots contained a greater proportion of forbs. However, for many phytophagous invertebrates forbs represent an important group of host plants. For this reason the absence of differences between the control (treatment 1) and those of the sown margins 8 and 9 in the majority of taxa were unexpected. However, the results presented here reflect only overall differences in abundance and species richness. In terms of species composition there were large differences in assemblage structure for most taxa, independent of the relative similarity in overall abundance and species richness. For example in the beetle fauna the sown treatments contained greater proportions of phytophagous species, reflecting the increased availability of forbs in treatments 8 and 9. Similarly for the spiders, the presence of statuesque plants such as thistles, kale or linseed, provided ideal web building structures for orb-web building spiders (e.g. Araneidae). The relative accessibility for higher trophic levels of the invertebrates within the sown and unsown treatments may also have differed, with many species within the grass based treatments being found at the bottom of the sward or within grass tussocks. While differences in both abundance and species richness in a number of groups were found between the sown treatments 8 and 9, these were relatively infrequent. This pattern is attributed to the rapid rise in floral dominance in many of the sown treatments by unsown species such as creeping thistle or nettles. This may have resulted in a homogenization in terms of floral composition between treatments 8 and 9, particularly as many of these unsown plant species support large phytophagous invertebrate faunas, e.g. thistles.

BirdsIn terms of providing foraging habitat for birds, there were clear benefits of some treatments, for some species, both in summer and winter. Considering first the grass treatments; in general, small insectivores, e.g. dunnock and granivorous finches and buntings e.g. Yellowhammer, made most frequent use of less intensively managed grass treatments (particularly uncut silage, treatment 7). However, the reverse was true for large insectivores e.g. Blackbird, which made more use of the intensively managed grass treatments, particularly in winter. These patterns reflect the abundance and accessibility of food resources, particularly invertebrate prey, supported by the different treatments. The abundance of the latter also generally increased with decreasing management intensity of the sward. Thus, the higher use by the smaller insectivores of treatment 7, and to a lesser extent 5 and 6, almost certainly represented a response to higher food abundance in these swards. In contrast, more frequent use of the more intensive treatments by large insectivores, despite the fact that most invertebrates including below ground species were less abundant in these treatments, almost certainly reflected the accessibility of invertebrate prey in these swards. Many of these species are known to prefer short swards for foraging but they

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will probably also require taller swards nearby as ‘sources’ of these prey, suggesting a mosaic of tall and short swards represents the optimum foraging habitat (Vickery et al 2001). Overall, these results provide evidence that more extensive management of grassland will deliver improved foraging opportunities for a number of grassland birds, as previously suggested by other work (e.g. Vickery et al. 2001, Atkinson et al. 2005). Areas of extensively managed land could be provided as whole or part fields and the cost effectiveness of different options needs to be assessed. The results also support previous findings that have highlighted the trade off, for grassland birds in particular, between providing swards that support abundant invertebrate prey and ones in which these prey are also accessible to birds (Devereux et al 2004, Atkinson et al 2005, Wilson et al 2005). The results of the present study suggests taller and denser swards can provide preferred foraging for small insectivorous species such as Robin and Dunnock and may act as ‘sources of invertebrates’ for larger insectivores that prefer to forage in short swards, suggesting the need for a mosaic of tall and short swards. The fact that so many insectivores used the sown treatments to a greater extent than the less intensively managed, more invertebrate rich, grass swards also supports the suggestion that accessibility of prey is a key determinant of habitat use by birds. It is possible that swards such as those created under treatment 7 might be made much more attractive and valuable to birds as a foraging habitat if they could be ’opened up’ by creating patches of short swards within them. This could be achieved through cutting, scarification or the use of graminicides, but the cost-benefits of such treatments would need to be assessed (e.g. Henderson et al 2007) as would the scale and configuration of such plots. The direct benefits of cutting, and the attractiveness of a short sward, was further suggested by foraging movements of birds as determined by radio tracking. Tagged blackbirds demonstrated an increased use of field margins for foraging after grass swards had been cut for silage in May. Considering the sown treatments, as predicted from previous work (e.g. Henderson et al 2004) these treatments were the most frequently used by granivores in winter, particularly the kale/quinoa mix in its first and second year. Undersown cereal was used much less frequently and use declined markedly after the first year. Interestingly, the kale/quinoa mix was also used more intensively than any of the grass treatments in winter and by small and large insectivores and, in general, they were also used more frequently than grass treatments in summer. The abundance of invertebrates was generally not significantly different in the sown treatments from that of control treatment 1, suggesting that the high use of sown treatments was not related to high prey abundance. We suggest, instead, that this is related to the increased accessibility of prey in these patchy sown swards compared with grass treatments. The value of these sown treatments for insectivores in summer was further supported by the fact that Dunnocks appeared to select hedgerows adjacent to sown treatments to nest. This species hold relatively small breeding territories in the summer and so the preference for nesting near these sown treatments probably reflects a preference for nesting near high quality foraging habitat. These results demonstrate that management options that provide margins, plots or even whole fields of seed-rich arable crops, will greatly enhance the foraging opportunities within grassland for both granivores and insectivores in winter and, to a lesser extent, in summer. The value of so-called ‘arable pockets’ has been suggested from analysis of large scale data sets (Robinson et al 2001) but this is the first time their value has been demonstrated in the field. The local extinction of many ‘arable‘ bird species has been attributed, in part, to the loss of foraging habitat in winter (Chamberlain & Fuller 2000) and over-winter survival is a key mechanism driving the declines of many granivores such as Yellowhammer (a species still found in grassland in good numbers) (Siriwardena et al 1998, 2000). Thus creating seed rich habitats within grassland may be a key recovery tool. It may also be possible to provide seed by allowing silage swards to set seed in winter. In this study there was extremely high use, particularly by Yellowhammers, of uncut silage (treatment 7) in its first winter, corresponding with the time when the rye grass was seeding. Whole field silage swards, dominated mainly by perennial ryegrass swards, left to set seed over winter are known to attract large numbers of buntings particularly Yellowhammers and Reed buntings (Buckingham & Peach 2006). The latter may provide a relatively easy way for a grassland enterprise to provide an increase in seed resources. However, a sward providing only grass seeds may be not prove as attractive to such a wide range of species as a wild bird cover crop or an undersown cereal, both of which provide a range of seeds over a longer period of the winter

Main implications InvertebratesPollinators: The grass-based treatments in this experiment were not readily manipulated to enhance the number or diversity of bumblebees. More interventionist measures, such as sowing flowers, are therefore needed to support bumblebees, but their effectiveness may be limited to the short-term. In contrast, grass-based plots, which are extensively managed, can enhance butterfly abundance and species richness. The introduction of novel habitats through sowing can also increase butterfly numbers and diversity, though with the information currently available, it is unclear how long this effect would be sustained.Beetles: The extensively managed grass-based treatments (5-7) supported high abundances and species richness of beetles, although there is evidence that margins requires 2-3 years to establish before these benefits are seen. The sown treatments were not found to be important for beetles.True bugs: Structurally complex treatments (treatments 5-9) supported high abundance and diversity of true bugs. Assemblage differences in grass and sown treatments drove the timing of abundance and diversity responses, with pioneer assemblages dominating sown plots within 1-2 years and grass seed specialists dominating grass plots by 2006.

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Plant hoppers: Plant hopper abundance increased with sward complexity, and particularly where sward height exceeded 5cm (treatments 3-7). However, species richness was unaffected by treatment with all grass-based treatments increasing in plant hopper species richness in each year. Sown treatments were characterised by low abundance and species richness of plant hoppers.Spiders: Spider abundance was linked with increased sward architectural complexity and so tended to be greatest in the extensively managed treatments, as well as treatment 4, which although cut twice yearly was cut at a lenient height.Orthoptera: There was no evidence of the margin management treatments, grass-based or sown, showing consistent across year effects on Orthoptera abundance. Margin management had no effect on dispersal.Flies: There was evidence that crane and St Marks fly abundance showed a preference for more structurally complex swards, such as those of treatments 5-9. Larvae: Larval abundance and biomass did not respond consistently to management treatments, however, in the grass-based treatment 2 abundance was lower in some years. Te response of larval mass and abundance to the grass-based treatments was generally weak.Slugs: There was no indication that any margin management treatments had consistent across year effect on slug abundance.

BirdsThe results show that grass swards managed under relatively extensive fertiliser cutting and grazing regimes, will provide enhanced foraging resources for a range of bird species throughout the year. Such swards support more abundant and diverse food resources in the form of both insects and seeds and are used more frequently by foraging birds than intensively managed grass swards. They could be provided as margins, part or whole fields and created by reducing fertiliser input as well as the cutting frequency and/or grazing intensity. The value of such grassland to most bird species is likely to be greatly enhanced if the relatively tall dense swards can be ‘opened up’ in a way that provides patches of short sward or bare ground. This will increase the accessibility of invertebrate prey to a range of insectivorous birds and extend the range of species able to benefit as well as the duration of that benefit. Further research is required to determine the optimal size and configuration of such patches for birds (determined by foraging efficiency and predation risk) and the most cost effective way of establishing such a patchy sward. Sown swards, particularly in the form of a wild bird cover crop but also as an undersown cereal, provide good foraging habitats for granivorous and insectivorous birds in winter, as do grass swards left to set seed in the first winter. Seed resources are rare in grassland in winter and even small areas of, for example wild bird cover crop, are likely to be used by relatively large numbers of birds throughout the winter.

References Atkinson, P.A., Fuller, R.J., Vickery, J.A., Conway. G., Tallowin, J.R., Smith, R., Haysom, K., Ings, T., Asteraki, E. & Brown, V.K. (2005) Influence of agricultural management, sward structure and food resources on grassland field use by birds in lowland England. Journal Applied Ecology 42, 932-942.Buckingham, D.L. & Peach, W.J. (2006) Leaving final cut grass silage in situ over winter as a seed resource for declining farmland birds. Biodiversity & Conservation, 15, 3827-3845Chamberlain, D.E. & Fuller, R.J. (2000) Local extinctions and changes in species richness of lowland farmland birds in England and Wales in relation to recent changes in agricultural land-use. Agriculture, Ecosystems and Environment 78, 1-17Devereux, C. L., McKeever, C.U. Benton, T.G. & Whittingham, M.J (2004). "The effect of sward height and drainage on Common Starlings Sturnus vulgaris and Northern Lapwings Vanellus vanellus foraging in grassland habitats." Ibis 146(s2): 115-122Ellenberg, H. (1988) Vegetation Ecology of Central Europe. Cambridge: Cambridge University PressGibson, C.W.D., Hambler, C., Brown, V.K., (1992) Changes in spider (Araneae) assemblages in relation to succession and grazing management. Journal of Applied Ecology 29, 132-142.Henderson, I.G., Vickery, J.A. & Carter, N. (2004) The use of winter crops by farmland birds in lowland England. Biological Conservation, 118, 21-32.Henderson, I.G., Morris, A.J., Westbury, D.B., Woodcock, B.A., Potts, S.G., Ramsay, A., Coombes, R. (2007) Effects Of Field Margin Management On Bird Distributions Around Cereal Fields. Aspects of Applied Biology 81: 53-60Hill M.O., Mountford J.O., Roy D.B., & Bunce R.G.H. (1999) Ellenberg's indicator valuesfor British plants. ECOFACT Volume 2 Technical Annexe. HMSO, NorwichHubbard C.E. (1972) Grasses. Penguin Books, UK.Robinson, R.A., Wilson, J.D., Crick, H.Q.P., (2001). The importance of arable habitat for farmland birds in grassland landscapes. Journal of Applied Ecology 38, 1059-1069.Siriwardena, G.M., Baillie, S.R., Buckland, S.T., Fewster, R.M., Marchant, J.H. & Wilson, J.D . (1998) Trends in abundance of farmland birds: a quantitative comparison of smoothed Common Birds Census indices. Journal of Applied Ecology, 35, 24-43.Siriwardena, G.M., Crick, H.Q.P., Baillie, S.R., & Wilson, J.D. (2000) Agricultural land use and the spatial distribution of granivorous lowland farmland birds. Ecography 23, 7002-719.

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Ter Braak, CJF and Smilauer, P (2002) Canoco Version 4.5, Biometrics-Plant Research International, WagenigenVickery, J.A, Tallowin, J.R., Feber, R.E., Asteraki, E.J., Atkinson, P.W., Fuller, R.J. & Brown, V.K . (2001) The management of lowland neutral grasslands in Britain: effects of agricultural practices on birds and their food resources. Journal of Applied Ecology, 38, 647-664.Williams, N.M., Minckley, R.L., Silveira, F.A., (2001) Variation in native bee faunas and its implications for detecting community change. Conservation Ecology 5, 57-89.Wilson, J. D., Whittingham, M. J., Bradbury, R. B. (2005). "The management of crop structure: a general approach to reversing the impacts of agricultural intensification on birds?" Ibis 147: 453-463.Woodcock, B.A., Potts, S.G., Pilgrim, E., Ramsay, A.J., Tscheulin, T., Parkinson, A., Smith, R.E.N., Gundrey, A.L., Brown, V.K., Tallowin, J.R., (2007) The potential of grass field margin management for enhancing beetle diversity in intensive livestock farms. Journal of Applied Ecology 44, 60-69.Woodcock, B.A., Potts, S.G., Westbury, D.B., Ramsay, A.J., Lambert, M., Harris, S.J., Brown, V.K., In press. The importance of sward architectural complexity in structuring predatory and phytophagous invertebrate assemblages. Ecological Entomology.

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References to published material9. This section should be used to record links (hypertext links where possible) or references to other

published material generated by, or relating to this project.Holt, C (2005) Managed lowland grassland and birds – the challenge of how to make intensively managed lowland grassland more interesting for birds. BTO News, Number 257, March-April 2005. Pp 20-21.

Pilgrim E.S.; Potts, S.G.; Vickery J.A.; Parkinson, A.E.; Woodcock B.A.; Holt, C.A.; Gundrey A.; Ramsay A.J.; Atkinson P.W.; Fuller R.J. and Tallowin, J.R.B. Enhancing wildlife in the margins of intensively managed grass fields. British Ecological Society Annual Meeting 5-7th September 2006, Oxford University

Potts, S.G. (2006) Enhancing the biodiversity value of intensive livestock farms: the PEBIL project. Case study for FAO report 'Status of World Pollinators' http://www.fao.org/AG/agp/agps/C-CAB/Castudies/pdf/6-020.pdf

Potts S.G (2005)., Invited keynote speaker, National Academy of Sciences (US) workshop on status of pollinators in North America, Washington DC, USA: Conservation of biodiversity of pollinators in natural and agro-ecosystems in Europe, 19 October 2005

Potts S.G (2005)., Invited keynote speaker, North American Pollinator Protection Campaign workshop, Washington DC, USA: Conservation of biodiversity of pollinators in natural and agro-ecosystems in Europe, 21 October 2005

Ramsay, A.J., Potts, S.G., Woodcock, B.A., Tscheulin, T., Brown, V.K., & Tallowin, J.R.B. (2005). Novel margin management to enhance biodiversity in intensive livestock farms. In: Proceedings of the British Society of Animal Science, pp. 66. BSAS, York, UK.

Ramsay, A.J., Potts, S.G., Woodcock, B.A., Brown, V.K. & Tallowin, J.B. (2004) Enhancing invertebrate biodiversity in intensive livestock farms using novel filed margin management. British Ecological Society Annual Meeting, 7-9 September 2004, Lancaster.   Ramsay, A.J., Potts, S.G., Woodcock, B.A., Tscheulin, T., Brown, V.K., & Tallowin, J.R.B. (2005) Novel margin management to enhance biodiversity in intensive livestock farms. In Proceedings of the British Society of Animal Science, pp. 66. BSAS, York, UK.

Ramsay A.J., Potts S.G., Woodcock B.A., Tscheulin T., Westbury D., Harris S., Tallowin J.R. & Brown V.K., Royal Entomological Society annual meeting, Sussex: Boosting bug diversity in an arable and pastoral landscape

Ramsay A.J. (2004) 'Marginally better? The effects of margins on grasshopper dispersal' at the Orthoptera SIG Conference, Royal Entomological Society, London 3rd November 2004.

Woodcock, B.A., Potts, S.G., Mortimer, S.R., Lawson, C.S., Ramsay, A.J., Brown, V.K., & Tallowin, J.R. (2005) The manipulation of field and field margin vegetation structure in intensively managed UK cattle grazed pasture systems: Implications for invertebrate biodiversity. In Proceedings of the British Society of Animal Science, pp. 231. BSAS, York, UK.

Woodcock B.A., Potts S.G., Ramsay A.J., Tscheulin T., Parkinson A., Smith R.E.N., Martyn T.M., Pilgrim E., Gundry A., Brown V.K. & Tallowin J.R. (2005 ), Royal Entomological Society annual meeting, Sussex: The potential for enhancing beetle diversity in intensive livestock farms using field margins

Woodcock B.A., Potts S.G., Ramsay A.J., Tscheulin T., Parkinson A., Smith R.E.N., Martyn T.M., Pilgrim E., Gundry A., Brown V.K. & Tallowin J.R. (2006), British Ecological Society annual meeting, Hertford: Enhancing invertebrate biodiversity in intensive livestock farms using novel field margin management

Woodcock B.A., Potts S.G., Mortimer S., Lawson L., Ramsay A.J., Brown V.K. & Tallowin J.R., (2005) British Society of Animal Science, York: The manipulation of vegetation field and field margin vegetation structure in intensively managed UK cattle grazed pasture systems: Implications for invertebrate biodiversity

Woodcock, B. A., Potts, S. G., Pilgrim, E., Ramsay, A. J., Tscheulin, T., Parkinson, A., Smith, R. E. N., Gundrey, A. L., Brown, V. K., and Tallowin, J. R. (2007). The potential of grass field margin management for enhancing beetle diversity in intensive livestock farms. Journal of Applied Ecology 44:60-69

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