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Botanical Assessment of Sown Grass Margins in the Countryside Stewardship Scheme

C N R Critchley, J A Fowbert & A J Sherwood1

Final Report

June 2005

ADAS Redesdale, Rochester, Otterburn, Newcastle upon Tyne NE19 1SB, UK 1ADAS Boxworth, Boxworth, Cambridge CB3 8NN, UK

MA01017 CSS Grass Margins ADAS June 2005

CONTENTS

Summary...... 4 Introduction...... 6 Methods...... 7 CSS prescriptions...... 7 Site selection ...... 7 Field survey...... 7 Data analysis...... 8 Results ...... 10 All margin types ...... 10 Sown grass margins ...... 16 Sown margins and environmental variables ...... 19 Older sown margins and environmental variables ...... 20 Rare species...... 22 Discussion ...... 23 Vegetation establishment...... 23 Vegetation development ...... 23 Variation in species composition ...... 23 Rare arable ...... 24 Methodological issues...... 24 Conclusions...... 25 Recommendations...... 25 Acknowledgements...... 26 References ...... 27 Appendices...... 31

3 MA01017 CSS Grass Margins ADAS June 2005 Summary 1. Field boundaries are important linear features in arable landscapes. The perennial grass or herbaceous verge next to the crop is beneficial to biodiversity and serves other functions but verges have been damaged and reduced in extent by intensive agricultural practices. To counteract this, the Countryside Stewardship Scheme (CSS) has encouraged farmers in England since 1993 to establish 6 m grass margins (management option ‘R3’) using either a limited selection of grass species or, more recently, a diverse mixture including perennial forbs. The CSS has made a substantial contribution to exceeding area targets set under the UK Biodiversity Action Plan for cereal field margins. 2. Most previous research on the establishment of grass margins has been short-term or applied at a limited number of sites, so the outcome of establishing grass margins at a national scale and over longer timescales was uncertain. The aim of this study was to assess the species composition of grass margins established in the CSS. 3. Four types of field margin were surveyed in 2004. These were grass margins sown with a simple grass seed mixture before 2002 (R3 pre-2002) or more recently (R3 post-2001), margins sown with a more diverse seed mixture including perennial grassland forbs (R3 + F) and normally cropped cereal headlands (controls). In total, a stratified random sample of 119 sites from eight Defra administrative regions was surveyed. At each site, botanical data were collected from 30 randomly located quadrats, soil cores were collected and the site searched for rare arable plant species. Data on management practices and habitat context were collected for each site. Data were analysed using ordination methods, Analysis of Variance and variation partitioning by Partial Redundancy Analysis. 4. In total, 264 taxa were recorded. Grass margins were markedly different from the control sites, which, apart from the obvious presence of the cereal crop and absence of other sown species, had lower abundance of grasses, forbs, perennials and larva and bumblebee foodplants. A small number of grass margins had been established by natural regeneration and tended to have more weedy perennials or annual arable species. The most abundant species at sown margins were the grasses included in the seed mixture. 5. A comparison of sown margins showed that R3 + F sites had higher species richness and numbers of sown species and perennial forbs than margins without additional forbs in the seed mixture. Forbs characteristic of grassland were only present at very low frequency in R3 pre-2002 and R3 post-2001 sites but, in contrast, were relatively common in R3 + F sites. Invasive, competitive perennials were, however, frequent at all sown sites. Annuals were more abundant in R3 post-2001 sites than older sown sites. 6. Most (85%) sown grass margin sites had loamy soil. Arable, woodland and grassland were the most common adjacent land uses and hedges, broadleaved trees and grass/tall herb verges the most common field boundary features. Two-thirds of sites over two years old were cut annually and more than half had cuttings removed. 7. The addition of forbs to the seed mixture had a significant effect on species composition of sown grass margins. Regions, soil extractable K, soil texture and age of site were also related to species composition. In sown margins older than two years, species composition was related to additional forbs in the seed mixture, regions, soil K and pH but not to age of margin. Region, soil properties and the addition of forbs tended to have independent (non-overlapping) effects on species composition. Species composition was not related to site management, adjacent land use, field boundary, slope, aspect or altitude. 8. No rare annual, arable plant species were found in CSS margins but there was one record of Apera spica-venti (loose silky-bent) at a control site. 9. In general, margins sown with simple grass seed mixtures were established successfully. Perennial grassland forbs only colonised at very low frequencies but did tend to establish and persist when included in the seed mixture. Therefore, their inclusion should be encouraged to enhance botanical diversity and foraging 4 MA01017 CSS Grass Margins ADAS June 2005 opportunities for nectar and pollen feeding invertebrates. The current mowing regimes are appropriate although more active management could be promoted to encourage further colonisation and spread of perennial grassland forbs. Grass margins should not be sited where annual plants are a priority for conservation.

5 MA01017 CSS Grass Margins ADAS June 2005 Introduction Field boundaries are important linear habitats in lowland arable farmland landscapes. The boundary usually comprises a hedge, trees, ditch, bank or other semi-natural feature but can also include a perennial grass or herbaceous verge next to the crop. The verge serves many functions potentially, including provision of valuable habitat for flora and fauna and a buffer zone that protects the remainder of the boundary from fertiliser and applied to the crop (Marshall & Moonen, 2002; Meek et al., 2002). The perennial vegetation also prevents annual weeds from establishing in the boundary and spreading to the crop (Smith et al., 1999b; Moonen & Marshall, 2001). However, in the UK and Western , many field boundaries have been removed as field sizes increased to accommodate modern farm machinery. In those remaining, the verge has often been lost or reduced in extent by mechanical cultivation close to the main field boundary feature. Contamination of verges by fertiliser and pesticides has also reduced their plant species diversity. To counteract these losses, research has focussed on methods of re-establishing perennial margins between the remaining field boundary and the crop. Experiments have shown that sown grass species can establish successfully. Species that form a dense, tussocky sward are advocated to provide habitat for invertebrates and other fauna (Thomas et al., 1991; Collins et al., 2003). Annual species are usually suppressed after the first year (Smith & Macdonald, 1989; Kirkham et al., 1999; Huusela-Veistola & Vasarainen, 2000; Asteraki et al., 2004; de Cauwer et al., 2004) and the risk of weeds spreading from new margins into the crop are low (Smith et al., 1999b). Grass margins have been established successfully in this way in an experimental agri-environment scheme (Critchley et al., 2004). The development of species-rich vegetation is, however, often inhibited by a paucity of colonisation sources and the low dispersal capability of many perennial grassland species (van Dorp et al., 1997; Kleijn et al., 1998). Establishment of a dense sward also presents few opportunities for additional species to regenerate. Experiments have also therefore focussed on sowing more diverse seed mixtures that include perennial forb1 species (Kirkham et al., 1999; Smith et al., 1999a; West et al., 1999; Asteraki et al., 2004; de Cauwer et al., 2004). These have been generally successful in establishing more species-rich perennial vegetation although the mowing regime can affect establishment rates (Smith et al., 1999a, de Cauwer et al., 2004). Margins with perennial forbs can increase arthropod diversity (Thomas & Marshall, 1999) and provide nectar and pollen food sources (Feber et al., 1996; Carvell et al., 2004). Most experiments have been short term (five years or less) although Bokenstrand et al. (2004) found that the majority of meadow species from a diverse seed mixture were still present after ten years. However, it is not known whether results from these experiments would be replicated if the methods were adopted by farmers at a national scale and over longer timescales. Since 1993, farmers in England, UK have been encouraged to establish grass margins under the Countryside Stewardship Scheme (CSS). In return for payments, farmers are required to establish and maintain a grass strip averaging at least 6 m wide (range 4 to 12 m) alongside a cropped field. Two establishment options are currently available to farmers, using either a limited selection of grass species from a defined list or a more diverse mixture that includes a range of grasses and forbs (Defra, 2003). By November 2004, 24,000 ha of grass margins had been established in this way under the CSS. This represents a substantial investment of resources and it is important therefore to determine whether the scheme has been successful in establishing the intended vegetation and maintaining it in the longer-term. The aims of this project were to assess the plant species composition of 6 m CSS grass margins of varying age and margins sown with additional forbs. Specifically, the following questions were addressed: 1) Does the intended perennial vegetation establish in grass margins? 2) Do perennial forbs colonise margins sown with simple grass seed mixtures? 3) Do sown perennial forbs establish and persist in grass margins? 4) Does species composition of grass margins vary with region, age, soil properties, management and habitat context?

1 A non-woody plant other than a grass, sedge or rush and in contrast to bryophytes, ferns and fern allies 6 MA01017 CSS Grass Margins ADAS June 2005 Methods

CSS prescriptions In the basic (R3) CSS grass margin option, farmers are required to sow at least six grass species that are suitable to the area and its soils from a defined list, at approximately 20 kg ha-1 with no single species comprising more than 20% (Appendix 1). For additional payment, a more diverse seed mixture including wildflower (forb) species can be sown (Options R3 + GS, GX or RS). Mesotrophic grassland species are most often incorporated in the seed mixture, for example Centaurea nigra (common knapweed), Achillea millefolium (yarrow), Leucanthemum vulgare (oxeye daisy), Prunella vulgaris (selfheal), Lotus corniculatus (bird’s-foot trefoil), Ranunculus acris (meadow buttercup), Plantago lanceolata (ribwort plantain) and acetosa (common ). The margin is then managed by cutting, usually after 15 July and uncut areas can be left each year. No agrochemical or fertiliser inputs are permitted except spot treatment to control pernicious weeds.

Site selection A site was defined as a single field margin of minimum length 120 m. Four types of margin were targeted for sampling. These were grass margins sown with a simple grass mix and more than two years old (R3 pre-2002) or more recently established (R3 post-2001), grass margins sown with a more diverse seed mixture including additional forbs (R3 + F) and normally cropped cereal field headlands as controls. The aim was to randomly select sites in clusters of four (one each of the four margin types) within a 10 x 10 km area and stratified by eight Defra regions (see Pywell et al., 2005b). The R3 + F sites were the least numerous and so these were selected first and clusters were based on their locations. There were too few R3 + F sites in some regions to draw the full sample so some clusters were based around other grass margins. Occasionally, a pre-2002 or post-2001 margin could not be located in a cluster. In total, 119 sites were surveyed (Table 1). A number of grass margins in the sample had been established by natural regeneration (three post-2001 and eight R3 pre-2002 margins).

Table 1. Number of sites surveyed in each region and management option. R3 = grass margins, R3 + F = grass margins with additional forbs. Region R3 pre-2002 R3 post-2001 R3 + F Control Total East 4 4 4 4 16 East Midland 3 4 0 4 11 North East 4 3 0 4 11 North West 4 4 4 4 16 South East 5 5 4 5 19 South West 4 4 4 4 16 West Midland 4 4 3 4 15 Yorks/Humber 4 4 3 4 15 Total 32 32 22 33 119

Field survey In each site, a 100 m long sampling zone was randomly located but not within 10 m of either end of the margin. The sampling zone encompassed the full width of the grass margin and was 6 m wide in control sites. In the zone, 30 nested quadrats of size 0.5 x 0.5 m were randomly located. To obtain top cover estimates, a single pinhit per quadrat was recorded by lowering a pin (diameter §PP YHUWLFDOO\LQWKHFRUQHURIWKHTXDGUDWDQGQRWLQJWKHILUVW hit of plant species, bare ground or litter. The presence of all additional plant species rooted in the quadrat (or bare ground or litter) was recorded cumulatively from the nested cells in the quadrat (12.5 × 12.5 cm; 25 × 25 cm; 50 × 50 cm). Species that could not be identified reliably in the field were recorded to genus level. Plants with less than two true were recorded as ‘seedlings’, and mosses and liverworts grouped as ‘bryophytes’. Plant species nomenclature is after Stace (1997).

7 MA01017 CSS Grass Margins ADAS June 2005 Each site was searched systematically for rare annual arable plant species on a target list compiled from a range of data sources (Wilson, 1999; Preston et al., 2002; Wilson & King, 2003) (Appendix 2), by walking in a zigzag pattern at a slow to moderate pace. The number of individuals per site of any rare species found was estimated in the field on a log scale. At the main locus of the rare species in a site, three 2 x 2 m quadrats were positioned 2 m apart and parallel to the field boundary. Plant species present in each quadrat were recorded and their percentage cover estimated. Grid references of rare species loci were recorded for future reference. Soil samples were collected from 3 randomly selected quadrats per site and from rare species loci. In sampling zones, ten evenly-distributed cores of depth 0-15 cm were collected per quadrat, and bulked to provide one sample per site. At rare species loci, ten cores were collected from the central quadrat (avoiding damage to the rare plants) and bulked. Soil samples were analysed by Direct Laboratory Services Ltd., Wolverhampton, using standard techniques (MAFF, 1986) for pH, extractable phosphorus (P), (K) and (Mg) and a hand texture assessment done. Texture classes were converted to estimated available water content (AWC) for data analysis (MAFF, 1988). Management information was collected for each site. Individual CSS agreements usually specified a list of species from which the farmer was required to sow a minimum number. Where possible, species actually sown were confirmed with the farmer. Species on the site list were assumed to have been sown if recorded in the field. The frequency (annual or less often) and timing (month) of cutting, treatment of cuttings (always removed, sometimes removed or not removed but pulverised, never removed) and presence of uncut areas in the sampling zone were recorded. Targeted weed control and grazing were also recorded. Data on the habitat context of each site were collected. Adjacent field boundary categories (grass/tall herb verge, hedge, broadleaved trees, coniferous trees, overhanging trees, ditch/open water), height estimates of woody species (if present), adjacent land cover beyond the field boundary (arable/horticulture/set-aside, grassland, woodland, built-up, open water, track), altitude (m), slope (sloping or level) and aspect (converted to absolute degrees from due south) were noted. Sites were surveyed during the period 7 June to 30 July 2004.

Data analysis All data were input from field forms to Microsoft Excel software and imported to Microsoft Access. All data were validated by comparing hardcopy outputs from Access with the original field forms.

Differences between margin types A Principle Components Analysis (PCA) was used to determine the overall variation in species composition at all 119 sites. A preliminary Detrended Correspondence Analysis showed a linear response of species along the first axis, confirming that PCA was the appropriate method to use (ter Braak & Šmilauer, 1998). This and subsequent multivariate analyses were carried out on log-transformed species frequencies (excluding unidentified seedlings and bryophytes) for each site, using Canoco V.4.02 software (ter Braak & Šmilauer, 1998). Plant cover was calculated as the percentage out of 30 pinhits per site and species frequencies as the number of quadrats per site in which they were recorded. Species numbers were the mean number per 0.25 m2 quadrat at a site. Differences between all field margin types were examined using one-way Analysis of Variance (ANOVA). Numbers and cover of all species and of species grouped according to and life history were analysed, as were pernicious weeds and foodplants of farmland birds, butterfly larvae and bumblebees. Bird and butterfly larva food plants were as listed in Smart et al. (2000) (Appendix 3). Bumblebee food plants were those identified in surveys of the Arable Stewardship Pilot Scheme (Pywell et al., 2005a; Appendix 3). Eight pernicious weeds were specified, namely Alopecurus myosuroides (black-grass), Anisantha sterilis (barren brome), Avena fatua (wild-oat), Cirsium arvense (creeping thistle), Elytrigia repens (common couch), Galium aparine (cleavers), (curled dock) and R. obtusifolius (broad-leaved dock). Cover values were transformed to arcsine x prior to analysis.

8 MA01017 CSS Grass Margins ADAS June 2005 Sixteen clusters in six regions had the complete set of sown CSS margin types (excluding naturally regenerated sites), i.e. one each of R3 pre-2002, R3 post-2001 and R3 + F. To determine differences between these margins and regional effects, a two-way ANOVA was carried out on mean regional values, with margin type and region as factors. Differences in soil properties between sown margin types and regions were also examined in this way, using the same set of sixteen clusters. Variation in soil properties amongst all field margin types was analysed using one-way ANOVA. ANOVAs were carried out using STATISTICA (Statsoft, Inc., 2001) software.

Species-environment relationships A variation partitioning procedure (Økland & Eilertsen, 1994; Økland, 2003) was used to determine the relative importance of different subsets of environmental variables in explaining variation in species composition of sown CSS margins. The full set of environmental data was available for 75 sown sites, comprising 24 R3 pre-2002, 29 R3 post-2001 and 22 R3 + F margins (naturally regenerated sites were excluded). Variation partitioning was performed using Redundancy Analysis (RDA) and Partial Redundancy Analysis (PRDA). Environmental variables were grouped into subsets (age, ± the addition of forbs, region, soil properties and habitat context). Site management was not included in this analysis because relatively few R3 post-2001 sites had a stable management regime. For each subset in turn, an RDA was carried out to select those variables that contributed significantly to the model (at P < 0.05) and to eliminate highly correlated environmental variables. Each variable was tested in turn using forward selection and Monte Carlo tests with 999 permutations. The significance of the first axis and the overall RDA were also tested using 999 Monte Carlo permutations. An RDA using the reduced (significant) sets of environmental variables was used to examine the overall relationship between species and the environmental factors. For each reduced environmental variable subset (i.e. including only the significant variables), the variation explained only by the subset was calculated from a PRDA in which the subset of interest was specified as environmental variables, and the remaining subsets as covariables. Variation shared by the subset with other variables was calculated as the total variation explained (TVE) by the subset of interest minus the variation explained only by the subset. This shared variation was further subdivided into all possible combinations of two or more variable subsets by carrying out a series of PRDAs in which the combined subsets of interest were environmental variables and the remainder were covariables. Variation partitioning produces a large number of variation components (2n – 1 where n is the number of variable subsets; potentially 63 in this case) and so to simplify the final results it is desirable to exclude those explaining negligible amounts of variation. A threshold for retention of these components was calculated as AVE = TVE/(2n – 1) where AVE is the average variation explained (Økland, 2003). Variation components that were less than AVE were rejected and the variation associated with them was redistributed equally among the components of order n – 1. For example, if the variation shared by age, region and soil properties was lower than the threshold value, the combination of age, region and soil properties was rejected and the associated shared variation was redistributed amongst age + region, age + soil properties, age + forbs, etc. The relationship between species composition of the older, established sites and environmental factors was then examined using the same variation partitioning procedure. In this case, site management was also included in the analysis. R3 pre-2002 margins and R3 +F margins more than two years old were analysed. There were 35 such sites available (23 R3 pre-2002 and 12 R3 + F margins, excluding naturally regenerated sites).

Phytosociological associations of rare species Data from the three quadrats at a rare species locus were used to identify the National Vegetation Classification (NVC) (Rodwell, 2000) plant community in which the rare species was growing. Similarity coefficients with NVC communities were calculated using both MATCH v2.16 (Malloch, 1999) and TABLEFIT v1.0 (Hill, 1996) software. Coefficients were used along with NVC floristic tables (Rodwell, 2000) to determine which community or sub- community the vegetation resembled most closely. 9 MA01017 CSS Grass Margins ADAS June 2005 Results

All margin types In total, 264 plant taxa were recorded (including crops but excluding unidentified seedlings and bryophytes). The PCA of all sites showed a clear separation along axis 1 of the control sites from R3 pre-2002 and R3 + F margins (Figure 1a). The R3 post-2001 sites showed some overlap in ordination space with the controls. Species were separated into five groups, albeit with some overlap (Figure 1b). These were cereal crops, annuals, sown grasses, sown forbs plus some sown grasses and ‘weedy’ perennials. Control sites coincided with crop species in the ordination space. Other margin types were separated less clearly but R3 post- 2001 tended to be more associated with annuals and R3 pre-2002 more with weedy perennials and sown species. Many R3 + F sites were associated with sown forbs; the exceptions were all younger sites. Some sites of all three CSS margin types were associated with the sown grasses. Naturally regenerated sites were associated either with weedy perennials or had species composition more similar to control sites. Many of the former had been regenerated from existing grassland or set-aside. The most widespread species at R3 pre-2002 sites were Dactylis glomerata (cock’s-foot), Phleum pratense (Timothy) and Festuca rubra (red fescue) (Table 2; see also Appendix 4). These grasses also had the highest mean within-site frequencies and were amongst those most commonly sown. D. glomerata was recorded from all R3 pre-2002 sites. At R3 post- 2001 sites, Lolium perenne (perennial ryegrass) was most widespread. L. perenne was sown at some sites (despite not being on the CSS list of recommended species) but its mean frequency was lower than frequencies for other sown grasses. Poa annua (annual meadow- grass), also widespread, is characteristic of early successional communities. The most widespread and frequent species at R3 + F sites were also perennial grasses, most of which would have been included in the seed mixtures. Centaurea nigra, a forb frequently sown at these sites, was also widespread. Other perennial grasses and forbs such as Agrostis stolonifera (creeping bent), Cirsium arvense and Ranunculus repens (creeping buttercup) had colonised widely in all three sown margin types, as had the annual Galium aparine. Triticum sp. (wheat) was, not surprisingly, most widespread and frequent in control sites; it was the sown crop at 19 (58%) sites but also occurred as a self-sown volunteer in others. Hordeum distichon (barley) was the sown crop at the remaining control sites, apart from one with a crop of Avena sativa (oat). Poa trivialis (rough meadow-grass) and Lolium perenne were amongst the most widespread species in all margin types, including control sites. There were significant differences between the main margin types in numbers or cover of most species groups (Table 3). The distinctiveness of the grass margins compared to the control sites was very evident (Plate 1). Apart from obvious differences due to the presence of a crop and absence of other sown species, the control sites had lower numbers and cover of grasses, forbs, perennials (especially perennial monocotyledonous species), and butterfly larva and bumblebee foodplants. Total cover of unsown species was also lower. Cover (but not numbers) of farmland bird foodplant species was higher in control sites, mainly because cereal crops are potential foodplants. Differences between the R3 post-2001 and other sites reflected their earlier successional state. R3 post-2001 sites had higher numbers and cover of annuals (particularly annual dicotyledonous species) and more plant litter cover.

10 MA01017 CSS Grass Margins ADAS June 2005 (a) (b)

(c) (d)

Plate 1. The four margin types: (a) simple grass margin more than 2 years old (R3 pre-2002), (b) recently established simple grass margin (R3 post-2001), (c) grass margin with additional forbs (R3 + F) and (d) normally cropped cereal headland (control).

11 MA01017 CSS Grass Margins ADAS June 2005 (a) +0.9 Axis 2

nr

nr

nr nr nr nr

nr nr

-0.5 nr nr -0.6 nr Axis 1 +0.9

(b)

ANNUALS

+0.5 Capsburs Poaannu Axis 2 Soncaspe Festprat Stelmedi Senevulg SOWN GRASSES Chenalbu Violarve Sperarve Polyavic Tripinod Sisyoffc Poaprat Festrubr Persmacu Matrdisc Phleprat Fallconv Cynocris VeropersLamipurp Lolipere Planmajo Tritsp. Phlebert Lotucorn Trifprat Dactglom Trisflav CEREAL CROPS Festarun Centnigr Rumeacet Leucvulg SOWN FORBS + GRASSES Horddist Trifdubi Agrocapi Trifrepe Galiveru Prunvulg Achimill Anthodor Planlanc Rhinmino Cirsarve Rubufrut Ranuacri Desccesp Rumeobtu Cerafont Agrostol Elytrepe Poatriv Heraspho WEEDY PERENNIALS Ranurepe Arrhelat

Holclana -0.4 -0.5 Axis 1 +0.7

Figure 1. Scatter plots of axes 1 and 2 from PCA ordination of all field margin types. (a) sites: R3 pre- 2002 (empty squares), R3 post-2001 (filled squares), R3 + F (crosses), controls (empty circles); naturally regenerated sites labelled ‘nr’. (b) species: only those with fit >7 shown for clarity; for species codes see Appendix 5.

12 MA01017 CSS Grass Margins ADAS June 2005 Table 2. The ten most common species in each sown margin type with % of sites in which they occur (mean % frequency in parentheses). Margin type codes as in Table 1.

R3 pre-2002 R3 post-2001 R3 + F Control n = 24 n = 29 n =22 n = 33

Dactylis glomerata 100 (72.4) Lolium perenne 75.9 (34.0) Festuca rubra 95.5 (82.9) Triticum sp. 78.8 (60.2) Phleum pratense 87.5 (51.3) Poa annua 75.9 (28.3) Dactylis glomerata 77.3 (33.2) Poa annua 57.6 (26.3) Festuca rubra 75.0 (46.3) Poa trivialis 72.4 (30.6) Poa trivialis 77.3 (31.1) Galium aparine 54.5 (13.5) Galium aparine 75.0 (8.5) Dactylis glomerata 69.0 (44.4) Lolium perenne 77.3 (25.0) Poa trivialis 48.5 (17.4) Agrostis stolonifera 70.8 (31.4) Phleum pratense 65.5 (40.5) Agrostis stolonifera 72.7 (28.0) Elytrigia repens 42.4 (7.4) Poa trivialis 70.8 (28.6) Festuca rubra 58.6 (44.7) Cynosurus cristatus 68.2 (39.8) Hordeum distichon 39.4 (38.8) Cirsium arvense 70.8 (15.0) Agrostis stolonifera 55.2 (10.0) Centaurea nigra 63.6 (22.7) Polygonum aviculare 39.4 (5.4) Ranunculus repens 62.5 (12.2) Galium aparine 55.2 (15.6) Ranunculus repens 63.6 (16.1) Lolium perenne 36.4 (8.5) 62.5 (8.2) Cirsium arvense 55.2 (1.5) Cirsium arvense 63.6 (11.5) Anisantha sterilis 27.3 (9.0) Lolium perenne 58.3 (22.1) Festuca pratensis 48.3 (28.7) Holcus lanatus 59.1 (19.8) dioica 27.3 (3.4)

13 MA01017 CSS Grass Margins ADAS June 2005 Table 3. Numbers and cover of species and species groups in the four main margin types with one-way ANOVA results. Data are means (±SE). Untransformed cover data are presented for clarity. Dicots = all dicotyledonous species, monocots = all monocotyledonous species; forbs excludes woody species. * P < 0.05; ** P < 0.01;*** < 0.001; ns = not significant.

R3 pre-2002 R3 post-2001 R3 + F Control F3,115 n 32 32 22 33

Numbers total 5.6 (0.22) 6.2 (0.43) 8.1 (0.62) 1.9 (0.23) 47.51 *** crops & volunteers 0.0 (0.02) 0.2 (0.28) 0.0 (0.03) 1.0 (0.10) 300.00 *** grasses 4.0 (0.20) 3.8 (0.30) 4.4 (0.27) 1.0 (0.12) 46.18 *** forbs 1.5 (0.13) 2.3 (0.34) 3.7 (0.56) 0.9 (0.17) 14.24 *** tree seedlings 0.1 (0.02) 0.1 (0.03) 0.1 (0.03) 0.0 (0.01) 1.70 ns horsetails & ferns 0.0 (0.03) 0.0 (0.02) 0.0 (0.01) 0.0 (0.01) 0.76 ns forb/grass ratio 0.40 (0.043) 1.35 (0.444) 0.93 (0.173) 1.30 (0.253) 2.50 ns annuals 0.5 (0.09) 1.8 (0.29) 1.1 (0.30) 0.8 (0.16) 6.66 *** perennials 5.0 (0.23) 3.9 (0.32) 6.8 (0.55) 0.7 (0.11) 68.42 *** annual dicots 0.3 (0.05) 1.5 (0.27) 0.9 (0.28) 0.6 (0.15) 6.64 *** annual monocots 0.2 (0.07) 0.3 (0.07) 0.2 (0.07) 0.2 (0.07) 0.75 ns perennial dicots 1.2 (0.12) 0.7 (0.13) 2.6 (0.47) 0.2 (0.04) 24.10 *** perennial monocots 3.8 (0.20) 3.2 (0.30) 4.1 (0.27) 0.5 (0.11) 53.75 *** weeds 0.7 (0.10) 0.6 (0.10) 0.4 (0.07) 0.4 (0.07) 3.22 * bird foodplants 2.5 (0.18) 3.3 (0.29) 3.4 (0.31) 1.9 (0.11) 10.54 *** bumblebee foodplants 1.0 (0.11) 1.2 (0.17) 2.8 (0.43) 0.4 (0.10) 20.84 *** butterfly foodplants 2.3 (0.12) 2.3 (0.20) 2.9 (0.33) 0.6 (0.07) 32.45 ***

Cover total vegetation 99.1 (0.48) 84.0 (4.14) 96.7 (2.61) 89.1 (3.06) 8.79 *** bare ground 0.3 (0.18) 9.3 (3.73) 2.7 (2.57) 9.1 (2.44) 5.80 ** litter 0.6 (0.38) 6.7 (2.53) 0.6 (0.46) 1.8 (1.41) 5.71 ** crops & volunteers 0 (0) 1.8 (0.59) 0 (0) 71.0 (4.35) 239.75 *** grasses 83.9 (2.97) 62.7 (5.64) 68.1 (5.65) 14.4 (3.49) 45.55 *** forbs 13.5 (2.47) 18.4 (4.03) 28.3 (4.80) 3.0 (0.82) 10.92 *** annuals 3.8 (1.74) 15.4 (3.54) 8.3 (3.14) 6.2 (2.08) 3.53 * perennials 94.8 (1.77) 61.5 (6.04) 87.1 (4.99) 7.0 (2.09) 96.18 *** weeds 11.5 (2.92) 9.2 (2.10) 5.4 (2.23) 5.0 (2.06) 3.31 * bird foodplants 37.5 (4.76) 36.6 (4.19) 45.7 (3.98) 78.2 (3.79) 24.39 *** bumblebee foodplants 10.4 (2.23) 9.5 (2.22) 23.8 (4.29) 1.9 (0.76) 14.32 *** butterfly foodplants 61.4 (3.56) 35.3 (4.25) 37.4 (5.15) 8.5 (3.00) 39.21 *** sown grasses & forbs 39.7 (5.96) 30.3 (6.19) 60.6 (6.34) 0 (0) 12.95 *** unsown species 59.2 (6.08) 51.3 (6.15) 36.1 (5.63) 17.4 (3.52) 27.91 ***

14 MA01017 CSS Grass Margins ADAS June 2005 The majority (85%) of sites were loamy soil, others being sands (9%) or clays (5%) (Table 4). No differences between margin types were detected for soil pH (F3,115 = 0.44, P = 0.72), extractable P (F3,115 = 0.17, P = 0.92) or K (F3,115 = 0.95, P = 0.42). However, extractable Mg was significantly higher in R3 pre-2002 margins (F3,115 = 3.24, P < 0.05). A further comparison by establishment method showed that sites naturally regenerated from existing grassland or set-aside tended to have relatively high levels of extractable Mg.

Table 4. Soil properties at sample sites (n = 119).

(a) Texture

Soil type Texture class No. of sites % of sites Loams Silty clay loam 36 30.3 Sandy silt loam 23 19.3 Silty loam 20 16.8 Sandy loam 15 12.6 Clay loam 4 3.4 Sandy clay loam 2 1.7 Organic sandy silt loam 1 0.8 Total 101 84.9 Sands Loamy sand 10 8.4 Organic loamy sand 1 0.8 Total 11 9.2 Clays Silty clay 6 5.0 Total 6 5.0 No data 1 0.8

(b) Chemical properties

Mean St. Dev. Min. Max. pH 6.8 0.79 5.0 8.3 Extractable P 28.5 14.89 5.0 95.0 Extractable K 207.5 118.30 44.0 722.0 Extractable Mg 122.2 91.61 14.0 542.0

15 MA01017 CSS Grass Margins ADAS June 2005 Sown grass margins In the comparison of sown margins, numbers of both species and perennial forbs per 0.25 m2 were significantly higher in R3 + F margins than R3 pre-2002 and R3 post-2001 grass margins (Table 5). The number of perennial forbs was more than 60% higher in R3 + F margins than R3 pre-2002 margins. The total number of sown species per 0.25 m2 was also substantially higher in R3 + F margins. However, there was no significant difference between margin types in the number of sown monocotyledonous species (grasses). Variation in the number of sown species present was therefore attributable to the additional forbs in the R3 + F seed mixtures. Forbs characteristic of perennial grassland were recorded relatively frequently in R3 + F margins, examples being Centaurea nigra, Achillea millefolium, Prunella vulgaris, Leucanthemum vulgare, Lotus corniculatus, Plantago lanceolata, Ranunculus acris, Rumex acetosa, Trifolium repens (white clover) and Galium verum (lady’s bedstraw) (see Appendix 4). These species tended to occur at equal proportions of R3 + F sites established before and after 2002, with the exception of Lotus corniculatus and Trifolium pratense (red clover), which were less prevalent at older sites. Most had also been recorded in R3 pre-2002 or R3 post-2001 margins but, in contrast, they were very sparse and often confined to just one or two records. R3 + F margins also had significantly more butterfly larva and bumblebee foodplant species per 0.25 m2 than the other grass margins. No differences were detected between margin types in cover or numbers of any of the unsown species groups. However, the total cover of unsown species was relatively high (c. 30-50%). Unsown perennial forbs that were recorded at relatively high frequencies in all three margin types were mostly invasive, competitive species such as Cirsium arvense, Rumex obtusifolius, (stinging nettle) and Taraxacum officinale (dandelion) (see Plate 2). Weed numbers did not differ significantly between margin types although there was a non-significant trend (P < 0.1) for lower weed numbers in R3 + F margins. The R3 post-2001 margins had significantly higher numbers and cover of annuals and cover of crop volunteers than the R3 pre-2002 and R3 + F margins. They also had less perennial cover. Numbers of annual monocotyledonous species differed between regions, being highest in the East. No other regional effects were detected. No differences were detected between sown margin types and soil properties. Soil pH varied significantly between regions (F5,10 = 8.67, P < 0.01), being highest in East region. Extractable K tended to be highest in South East and South West regions; this was just outside the limits of statistical significance (F5,10 = 2.81, P = 0.08).

Plate 2. Pre-2002 grass margin colonised by Cirsium spp. (thistles).

16 MA01017 CSS Grass Margins ADAS June 2005 Table 5. Numbers and cover of species and species groups in sown margins in six regions with two-way ANOVA results. Data are means (±SE). Cover data are backtransformed; standard errors have two (asymmetrical) values, which are + and – respectively. Monocots = all monocotyledonous species; forbs excludes woody species. * P < 0.05; ** P < 0.01;*** < 0.001; ns = not significant. (a) Numbers

Margin Type Region ANOVA Variable R3 pre- R3 post- R3 + F E NW SE SWWM YH Type Region 2002 2001 n 6 6 6 3 3 3 3 3 3 F2, 10 F5, 10

Total 5.9 (0.55) 6.2 (0.71) 7.9 (0.65) 6.9 (1.04) 7.1 (0.69) 6.5 (0.26) 8.4 (0.87) 4.7 (0.77) 6.5 (1.38) 4.75* 2.86 Annual forbs 0.3 (0.07) 1.6 (0.42) 0.6 (0.20) 0.5 (0.12) 1.3 (0.60) 0.4 (0.20) 1.4 (0.97) 0.5 (0.37) 0.9 (0.15) 6.29* 1.35* Annual monocots 0.2 (0.07) 0.3 (0.09) 0.1 (0.07) 0.5 (0.02) 0.1 (0.02) 0.2 (0.18) 0.1 (0.08) 0.1 (0.04) 0.1 (0.09) 2.22 3.45* Perennial forbs 5.8 (0.98) 3.8 (0.79) 9.5 (0.93) 6.3 (2.77) 6.0 (1.15) 6.8 (2.42) 6.7 (1.74) 5.0 (0.58) 7.5 (2.93) 7.73** 0.34 Perennial monocots 4.0 (0.42) 3.4 (0.53) 4.4 (0.40) 4.4 (0.25) 4.3 (0.31) 4.1 (0.50) 4.9 (0.81) 2.8 (0.57) 3.2 (0.81) 1.66 2.06

Total unsown 3.6 (0.39) 4.0 (0.67) 3.0 (0.29) 2.6 (0.13) 3.8 (0.89) 3.9 (0.56) 4.7 (0.96) 2.6 (0.48) 3.4 (0.30) 1.34 1.69 Unsown perennial 1.0 (0.17) 0.5 (0.11) 0.8 (0.16) 0.6 (0.13) 0.8 (0.17) 1.1 (0.38) 0.8 (0.19) 0.7 (0.12) 0.9 (0.36) 2.22 0.59 forbs Unsown monocots 2.1 (0.31) 1.7 (0.13) 1.5 (0.22) 1.5 (0.09) 1.6 (0.57) 2.0 (0.30) 2.4 (0.46) 1.5 (0.05) 1.6 (0.11) 2.06 1.39

Total sown 2.3 (0.37) 2.2 (0.52) 4.9 (0.64) 4.2 (0.92) 3.3 (1.03) 2.6 (0.74) 3.7 (1.54) 2.0 (1.06) 3.1 (1.15) 9.26** 1.19 Sown monocots 2.2 (0.40) 2.2 (0.52) 3.1 (0.49) 3.4 (0.12) 3.1 (0.79) 2.4 (0.85) 2.8 (0.71) 1.5 (0.53) 1.9 (0.63) 1.15 1.24

Bird foodplants 2.6 (0.29) 3.5 (0.48) 3.4 (0.20) 3.1 (0.32) 3.8 (0.70) 3.3 (0.18) 3.6 (0.63) 2.3 (0.46) 2.9 (0.51) 2.05 1.30 Butterfly foodplants 2.3 (0.13) 2.2 (0.33) 3.2 (0.30) 2.6 (0.32) 3.2 (0.20) 2.4 (0.14) 3.0 (0.68) 1.9 (0.19) 2.3 (0.72) 5.27* 1.87 Bumblebee foodplants 1.1 (0.23) 1.4 (0.26) 2.8 (0.44) 1.3 (0.65) 1.5 (0.30) 1.4 (0.50) 2.2 (0.92) 1.2 (0.38) 2.3 (1.05) 9.21** 1.14 Weeds 0.5 (0.09) 0.5 (0.07) 0.3 (0.06) 0.5 (0.04) 0.5 (0.17) 0.5 (0.12) 0.3 (0.11) 0.4 (0.04) 0.5 (0.19) 3.58 1.00

17 MA01017 CSS Grass Margins ADAS June 2005 (b) Cover

Margin Type Region ANOVA Variable R3 pre-2002 R3 post-2001 R3 + F E NW SE SWWM YH Type Region n 6 6 6 3 3 3 3 3 3 F2, 10 F5, 10

Total 99.4 90.1 98.6 99.0 92.0 97.5 96.7 99.6 94.6 2.43 0.49 (0.49, 0.94) (4.96, 6.34) (1.23, 2.42) (1.05, 3.10) (5.96, 9.41) (1.90, 3.12) (2.56, 4.27) (0.37, 1.12) (5.37, 14.96) Crops 0 1.3 0 0 0 0.2 0.2 0.2 0.8 7.47* 1.00 (0, 0) (1.08, 0.75) (0, 0) (0, 0) (0, 0) (0.56, 0.19) (0.56, 0.19) (0.74, 0.25) (2.32, 0.78) Annuals 1.3 21.4 2.4 11.6 9.5 2.0 6.2 4.8 5.2 8.68** 0.58 (1.80, 1.05) (4.80, 4.44) (2.59, 1.66) (1.33, 1.27) (7.36, 5.42) (5.70, 1.95) (16.95, 6.15) (7.01, 3.98) (6.80, 4.05) Perennials 95.9 57.9 93.9 85.0 78.3 88.0 86.0 92.3 83.8 9.73** 0.28 (1.88, 2.43) (7.98, 8.19) (3.61, 5.08) (3.21, 3.52) (13.56, 17.83) (5.16, 6.37) (12.56, 22.25) (6.58, 11.74) (14.71, 26.00) Monocots 84.4 62.6 70.2 80.2 69.1 69.5 71.6 79.5 66.6 2.18 0.29 (5.34, 6.22) (9.32, 9.82) (2.77, 2.84) (6.21, 7.05) (13.73, 15.72) (9.21, 10.07) (15.49, 18.72) (4.00, 4.32) (13.42, 14.96) Forbs 10.9 19.3 25.0 13.8 19.7 22.6 18.6 17.7 16.1 1.03 0.09 (5.56, 4.53) (8.51, 7.27) (3.37, 3.23) (11.18, 8.24) (9.82, 8.24) (5.47, 5.04) (20.83, 14.14) (1.92, 1.84) (9.06, 7.35) Total unsown 48.3 52.5 30.3 39.4 41.4 48.4 42.4 56.0 34.1 2.35 0.48 (7.61, 7.57) (8.34, 8.41) (4.79, 4.58) (4.11, 4.03) (15.79, 14.93) (13.21, 13.09) (20.30, 19.04) (6.95, 7.07) (2.31, 2.27)

18 MA01017 CSS Grass Margins ADAS June 2005 Sown margins and environmental variables At the sub-sample of sown margins, the most common adjacent land use was arable, followed by woodland and grassland (Table 6). Hedges were present in the field boundary at more than half of the sites, and broadleaved trees and grass/tall herb verge also occurred widely. Mean altitude was 97 m above sea level (range –4 to 160 m) and median field boundary height category was 2-4 m.

Table 6. Incidence of land cover and field boundary categories adjacent to sown margins (n = 75). Sites can have more than one category present. Arable includes horticulture and set-aside; built-up is roads and buildings.

Adjacent land cover No. of sites Field boundary No. of sites Arable 42 Hedge 41 Woodland 12 Broadleaved trees 29 Grassland 11 Grass/tall herb verge 27 Built-up 6 Overhanging trees 9 Open water 2 Ditch/open water 9 Track 1 Coniferous trees 4

The addition of forbs to the seed mixture and the age of margin both had a significant relationship with species composition (Table 7). Three regions (South West, East and East Midlands) and two soil variables (extractable K and soil AWC) were also significant. However, species composition was not significantly related to any of the habitat context variables. Regions explained the most variation, followed by soil, margin type (± forbs) and age. However, the addition of forbs was the strongest single explanatory variable. The threshold for retention of variation components was 1.25%. None of the second or higher order partial intersections explained more than this even after the redistribution procedure. Therefore, none was included in the final model. This indicated that there was very little overlap in the variation explained by margin type, age, region or soil properties, i.e. these environmental variable subsets acted independently of one another in explaining variation in species composition. The largest shared variation component was of region and soil properties, but even this only explained 0.6%.

Table 7. Variation explained by each environmental variable subset from RDAs of sown grass margins. TVE = total variation explained by all significant variables. See Figure 2 for variable codes.

Subset Significant Variation explained % of TVE Axis 1 (F) Overall (F) variables (%) Option ± forbs 4.5 24.5 n/a 3.44**

Age age 3.1 16.8 n/a 2.32**

Region SW 7.8 42.4 2.35** 2.01** E EM

Soil K 4.4 23.9 1.96** 1.67** AWC

All sig. vars 18.4 3.78** 2.15**

The strong effect of additional forbs in the seed mixture was shown by its correlation with axis 1 in the biplot of species and the significant environmental variables (Figure 2). There was a clear tendency for perennial forbs to be present at sites where they were included in the seed mixture. Soil AWC was also correlated with axis 1; Ranunculus repens and R. acris tended to occur on soils with higher water capacity. Site age and soil extractable K were correlated with axis 2. Naturally colonising perennial species such as Agrostis stolonifera, Rumex obtusifolius and Elytrigia repens were associated with older sites whereas annuals 19 MA01017 CSS Grass Margins ADAS June 2005 (e.g. Sisymbrium officinale (hedge mustard), Veronica persica (common field speedwell) and Capsella bursa-pastoris (shepherd’s purse)) were associated with younger sites. Some sown grasses (Phleum pratense and Poa pratensis (smooth meadow-grass)) were associated with younger sites, indicating that they established at an early stage. The South West region was separated from East and East Midlands regions along axis 3, indicating some differences in species composition between these regions. For example, Alopecurus myosuroides (black- grass) and Elytrigia repens were less likely to occur in the South West.

Age +0.8 Axis 2

Pteraqui Holclana Agrostol Elytrepe Vicitetr

AWC Ranurepe Ranuacri Anthodor PlanlancDauccaro F Achimill Trifprat Galiveru Prunvulg Festrubr E Fagusylv Rumeacet Centnigr Cynocris Lotucorn Stelmedi Planmedi Siledioi IrisfoetMedilupuTilix eu Hiersp. Leucvulg Galimoll Malvmosc Picrechi Festovin Chenalbu Corosqua Brasoler Dipsfull Phlesp. Poaannu Pisusati Geradiss Poaprat EM Veropers Cardsp. Veroserp Raphraph SW Capsburs Soncaspe Soncoler

K -0.7 -0.8 Axis 1 +1.0

Figure 2. Biplot of axes 1 and 2 from RDA of sown margins. AWC = soil estimated available water content, E = East region, EM = East Midlands region, F = added forbs in seed mixture, K = soil extractable K, SW = South West Region. Only species with fit >9 shown for clarity; for species codes see Appendix 5.

Older sown margins and environmental variables Sites ranged from 3 – 15 years old, though all but three were within the range 3 – 7 years. Approximately two-thirds of the sites were cut annually and all except three were cut in July or later (Table 8). One third of sites had not been cut recently and almost half (43%) had some uncut areas in the sampling zone at the time of survey. More than half of the remainder always had the cuttings removed. Targeted weed control was carried out at more than half of the sites but only three had ever been grazed. The addition of sown forbs had a significant relationship with species composition at the older sown sites, as did three regions (South West, East and Yorks/Humber), soil extractable K and soil pH (Table 9). However, none of the habitat context variables, management variables or age of site were related to species composition. Collectively, the three regions explained the greatest percentage variation but the addition of sown forbs was the single most important variable.

20 MA01017 CSS Grass Margins ADAS June 2005 Table 8. Management applied at sown margins greater than two years old. (n = 35; data incomplete for some sites).

No of No of sites sites Cutting: frequency annual 21 Cuttings removed always 13 less often 12 sometimes or pulverised 7 never 11 Cutting: month before July 3 July 16 Weed control 19 after July 11 Grazing 3 Uncut areas present 15

Table 9. Variation explained by each environmental variable subset from RDAs of sown grass margins greater than two years old. TVE = total variation explained by all significant variables. See Figure 3 for variable codes. Note: variation explained cannot be contrasted with Table 7 because these are relative values not comparable between different datasets (Økland, 1999).

Subset Significant Variation explained (%) % of TVE Axis 1 (F) Trace (F) variables Option ± forbs 7.7 28.0 n/a 2.75, 0.001

Region SW 14.6 53.1 2.41** 1.76** E YH

Soil K 9.5 34.5 1.81* 1.68** pH

All sig. vars as above 27.5 3.09** 1.77**

The retention threshold for the variation partitioning procedure was 3.6%. However, as with the previous analysis, none of the partial intersections (components of variation that were shared between two or more environmental subsets) explained more than 3.6%. The largest shared component after redistribution was 2.7%, which was between region and soil properties. As with the larger subsample of sown margins therefore, the added forbs, region and soil properties tended to act independently of one another in explaining variation in species composition. The sown grasses Dactylis glomerata and Phleum pratense were clearly related to the R3 pre-2002 margins, whereas a notable number of perennial forbs were associated with the R3 + F margins (Figure 3). The Yorks/Humber region was separated from East and South West regions in the ordination space; the latter were associated with higher soil pH values. The commonly sown Phleum bertolonii (smaller cat’s-tail) and Poa pratensis, along with annuals such as Anisantha sterilis and Veronica arvensis (wall speedwell) were also associated with the latter.

21 MA01017 CSS Grass Margins ADAS June 2005 pH Poaprat +0.7

Axis 2 Phlebert Veroarve SW Dactglom E Anisster Soncoler Festovin Silelati K Festrubr Malvmosc Planmedi Hiersp. Galimoll Phleprat Tilix eu R3 Paparhoe Raphraph Veroserp Sisyoffc Cynocris Irisfoet Veropers Trisflav Siledioi Leucvulg Fagusylv Lapscomm Rumeacet Elytrepe Dauccaro Trifprat Centnigr Prunvulg F Galiveru Rumeobtu Trifcamp

Achimill YH Planlanc Ranuacri Ranurepe Poatriv -0.6 -0.8 Axis 1 +1.0

Figure 3. Biplot of axes 1 and 2 from RDA of sown margins greater than two years old. E = East region, F = R3 + F, K = soil extractable K, R3 = R3 pre-2002, SW = South West region, YH = Yorks/Humber region. Only species with fit >14 shown for clarity; for species codes see Appendix 5.

Rare species No rare arable species from the target list were found at any of the CSS grass margin sites. One population (>1000 individuals) of Apera spica-venti was found at a control site of wheat in East Midlands region. There are historical records of the species in the same 10 x 10 km square during 1970-1986 and in an adjacent square during 1987-1999 (Preston et al., 2002). There were only six companion species recorded in the plant community (Artemisia vulgaris (mugwort), Poa annua, Silene latifolia (white campion), Geranium robertianum (herb robert) and G. pusillum (small-flowered crane’s-bill)), all at less than 1% cover in each quadrat. Because of this, the community did not closely resemble any NVC classes but the closest matches were to OV20 (Poa annua – Sagina procumbens) and OV21 (Poa annua – Plantago major). Details of this record are in Appendix 6.

22 MA01017 CSS Grass Margins ADAS June 2005 Discussion

Vegetation establishment The vegetation of grass margins contrasted strongly with that of normally cropped control sites, being composed primarily of perennial species. In recently established grass margins however, the annual component of the vegetation was similar to the annual weed community present in the cropped sites. Even in these younger sites, the sown grasses had already established and were prominent in the vegetation. By the third year, annuals had declined substantially and perennial species were dominant. This reflects the early successional sequence at experimental sites (Smith & Macdonald, 1989; Kirkham et al., 1999; Asteraki et al., 2004; de Cauwer et al., 2004) and on set-aside land sown with green cover (Firbank et al., 2003). After two years, age had no further detectable effect on the species composition of the CSS sown grass margins. The vegetation of naturally regenerated sites reflected their origins, having an annual weed component at former arable sites and more ruderal or grassland species at former set-aside or grassland sites.

Vegetation development Perennial grassland forbs did establish from the seed mixture in R3 + F sites. Perennial grasses were still the main component at these sites but the presence of grassland forbs distinguished them from the other sown grass margins, where these forbs were largely absent. The grassland forbs in the R3 + F sites almost certainly originated from the seed mixture. This concurs with findings from small-scale experiments and confirms the benefits of sowing the more diverse seed mixtures. In sites older than two years, there was no detectable effect of age on species composition. This indicates that the grassland forbs persisted in the older R3 + F sites although the legumes Lotus corniculatus and Trifolium pratense were less frequent in older sites. It also suggests that there had been no progressive colonisation at R3 pre-2002 margins. Competitive or ruderal species colonised both margin types but did so at an early stage. Therefore, stable plant communities had established at a relatively early stage from both diverse and simple seed mixtures. An advantage of establishing a competitive sward at an early stage is that the risk of colonisation by weed species is reduced (Smith et al., 1999b). The inclusion of less competitive species in the more diverse seed mixture could have increased the risk of weed invasion but this did not occur. Although spot treatment of weeds was carried out at many sites, this was done equally at R3 pre-2002 and R3 + F sites. Overall, the evidence suggests that there was little difference in the invasibility of plant communities from simple and diverse seed mixtures. There are conflicting views about the relationship between species richness and community invasibility. Reducing the plant species richness of grassland communities can increase their vulnerability to invasions (Knops et al., 1999). Conversely, the presence of competitive species was shown to be more important than species richness in determining the rate of species invasions in productive grassland (Crawley et al., 1999). In CSS sown margins the presence of competitive grasses appeared to be more important. Competitive grasses were present at high frequencies in both R3 pre- 2002 and R3 + F margins but there was no suggestion that invasibility might be related to species richness. As well as increasing plant species diversity, the addition of sown forbs increased potential foraging species for both butterfly larvae and bumblebees. All grass margins had more foodplant species than the cropped control sites. The presence of foodplant species does not necessarily confer advantages to these groups unless they are available, utilised and limiting to their populations. However, many species have declined in intensive agricultural landscapes and additional nectar and pollen sources are readily utilised (Pywell et al., 2005a). Compared to control sites, grass margins also had more bird foodplant species but this was countered by the absence of the cereal crop, which is also a potential food source.

Variation in species composition Differences in species composition between sown grass margins were largely due to the addition or otherwise of perennial forbs to the seed mixture. There was a relatively large amount of unexplained variation in the multivariate analyses. This is partly an artefact of the

23 MA01017 CSS Grass Margins ADAS June 2005 analytical method itself (Økland, 1999) but it is likely that the actual composition of the seed mixture, which varied between sites, would account for much of the unexplained variation, since most sown species did appear to establish. Some grass species were sown almost universally and this would reduce the variation in species composition between sites. In the CSS, farmers were encouraged to sow species appropriate to the geographic location and local conditions. This might account for some of the variation between regions. However, the higher number of annual monocotyledons found in East region does concur with previous surveys, which showed that in intensive arable landscapes (including the East region) species of early successional stages tend to be more prominent than in mixed farming landscapes (Critchley & Fowbert, 2000; Critchley et al., 2004). The effect of region in explaining variation in species composition was independent of soil properties and habitat context. This is surprising, since soil properties did vary between regions, and the amount of semi-natural habitat also differs between intensive arable and mixed farming regions. This suggests that other factors operate at a regional scale, for example as a consequence of differences in climate and local farming practices. Habitat context is more likely to influence naturally regenerated sites because they are more susceptible to colonisation from adjacent habitats. Habitat context was not related to species composition of CSS sown margins. This contrasts with studies of field boundary ground layer vegetation (Le Coeur et al., 1997) and a range of recently sown field margins (Critchley et al., 2004) that showed some variation in plant species composition was related to local landscape structure. In the CSS grass margins, the sown grasses established quickly and a closed sward developed. Adjacent habitat is less likely to influence species composition if there are few opportunities for seedling establishment. Cutting regime or other management practices were not related to species composition. This was surprising because previous research showed that the cutting regime affected diversity, particularly of forbs (Smith et al., 1999a; Schippers & Wouter, 2002; de Cauwer et al., 2004). However, the management at most CSS sites was conducive to maintaining diversity, as the majority were cut annually and more than half had cuttings removed. In this national sample, the relatively small differences in management practices might not have resulted in major differences in species composition. In a closed sward, more severe disturbance might be needed before additional species can colonise (Burke & Grime, 1996). However, differences in vegetation structure arising, for example, from uncut areas, will increase diversity of some invertebrate groups (Mitchley, 1994).

Rare arable plants No rare annual plant species were found in grass margins but this was not unexpected. The dense, closed grassy sward would offer few opportunities for these species to germinate and complete their life cycle, although some populations might survive at the edge of the grass margin next to the crop. Field margin management that includes regular cultivation and produces open vegetation is more suitable for annual plants and rare species have been recorded there in agri-environment schemes (Critchley et al., 2001; 2005). In normally managed cereal crops, rare species might germinate but fail to complete their life cycle, so depleting the population.

Methodological issues A list of species included in the seed mixture at each site was obtained but some discrepancies can be expected between these lists and the exact species sown. For example, records held by farmers might not be complete or some contamination of seed mixtures with other species might occur. The assumption was made that if a species was recorded at a site where it was sown, then it had originated from the seed mixture. It is possible that some species, especially common perennial grasses, might regenerate naturally as well as from the seed mixture. In these cases, the incidence of unsown species would have been underestimated. However, since perennial grassland forbs were virtually absent from R3 pre- 2002 and R3 post-2001 sites, it can be safely assumed that they were unlikely to colonise naturally at R3 + F sites either, so that any differences in this group of species would be attributable to the seed mixture.

24 MA01017 CSS Grass Margins ADAS June 2005 It is likely that some species occurring at low frequencies would not be recorded in quadrats. Some sites had been mown before the field visits. Plant species were identified from vegetative parts and, although some species might not have been so noticeable in the sward, this was not thought to have affected accuracy of records.

Conclusions Addressing the initial questions: 1) The intended perennial vegetation did establish in the sown grass margins. The grass margins contrasted sharply with the cropped control sites and the successional process reported from previous small-scale experiments was reflected across this national sample. 2) Perennial forbs associated with semi-natural habitats did not colonise grass margins sown with simple grass mixtures. Competitive and ruderal forbs did, however, colonise grass margins of all types. 3) Sown perennial forbs did establish in R3 + F margins, and they persisted in the older margins. 4) Species composition was related to site age, but not in sites over two years old. Soil properties and region had some relationship with species composition, but they tended to act independently. Management and habitat context had no detectable relationship with species composition. The UK Biodiversity Steering Group Report (Anon., 1995) has set a target to restore the biodiversity of 15,000 ha of cereal field margins by 2010, to which the CSS grass margins make a substantial contribution. It is important though to consider the quality of the resource provided as well as the extent. The inclusion of additional forbs in seed mixtures was successful in the CSS and increased the biodiversity value of these sites. Sites sown with simple grass mixtures had limited botanical diversity but should provide benefits to invertebrates and act as a buffer zone between the crop and the remainder of the field boundary. Under the CSS at this national scale, farmers established and maintained grass margins successfully. The absence of rare arable plant species from the CSS sample and the decline in annuals after the first two years shows that grass margins are unsuitable for the conservation of rare annual arable plants. It will be important, therefore to target grass margins away from sites where they are likely to occur.

Recommendations 1. Grass margins sown with simple grass seed mixtures (i.e. six or more common grass species) should be recommended where a rapid, reliable and relatively cheap method is needed to create a buffer zone between the crop and field boundary, or a dense, grassy habitat. 2. Grass margins are relatively unaffected by habitat context and so their establishment could be encouraged at difficult sites, for example where there are overhanging trees. They could also be sited next to valuable habitats if a buffer between the habitat and crop is desirable. 3. The addition of perennial grassland forbs to the seed mixture should be encouraged to increase botanical diversity and foraging potential for nectar and pollen feeding invertebrates. Most of the forbs widely included in the current seed mixtures are appropriate. 4. Grass margins should not be sited where rare annual arable plants are likely to be present, nor where the priority is to conserve annual plant communities. 5. The current management regime of regular cutting in late summer, with removal of cuttings wherever possible, can be continued. More active management to promote colonisation and spread of desirable forbs should also be encouraged.

25 MA01017 CSS Grass Margins ADAS June 2005 Acknowledgements This project was funded by the Department for Environment, Food and Rural Affairs. We are grateful to farmers for allowing access to their land, to Mark Stevenson and Simon Smith of Defra for providing background information and to Richard Pywell and Mark Stevenson for site selection. The contract was managed by Andy Parkin and field data were collected by Maurice Bailey, Ben Benatt, Hugh Dixon, Ruth Hadden, John Larbalestier, Caroline Munns, Tony Smith and James Towers.

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27 MA01017 CSS Grass Margins ADAS June 2005 Kirkham, F.W., Sherwood, A.J., Oakley, J.N. & Fielder, A.G. (1999). Botanical composition and invertebrate populations in sown grass and wildflower margins. Field margins and buffer zones: Ecology, Management and Policy, (eds N. D. Boatman, D. H. K. Davies, K. Chaney, R. Feber, G. R. de Snoo, and T. H. Sparks) Aspects of Applied Biology 54, 291-298. Kleijn, D., Joenje, W., Le Coeur, D. & Marshall, E.J.P. (1998). Similarities in vegetation development of newly established herbaceous strips along contrasting European field boundaries. Agriculture, Ecosystems & Environment, 68, 13-26. Knops, J.M.H., Tilman, D., Haddad, N.M., Naeem, S., Mitchell, C.E., Haarstad, J., Ritchie, M.E., Howe, K.M., Reich, P.B., Siemann, E. & Groth, J. (1999). Effects of plant species richness on invasion dynamics, disease outbreaks, insect abundances and diversity. Ecology Letters, 2, 286-293. Le Coeur, D., Baudry, J. & Burel, F. (1997). Field margins plant assemblages: variation partitioning between local and landscape factors. Landscape and Urban Planning, 37, 57-71. MAFF (1986). The Analysis of Agricultural Materials, MAFF Reference Book 427, 3rd edition. HMSO, London. MAFF (1988). Agricultural Land Classification of England and Wales. Revised Guidelines and Criteria for Grading the Quality of Agricultural Land. MAFF, Alnwick. Malloch, A.J. (1999). MATCH version 2 for Windows NT/95/98/2000. University of Lancaster, Lancaster. Marshall, E.J.P. & Moonen, A.C. (2002). Field margins in northern Europe: their functions and interactions with agriculture. Agriculture, Ecosystems & Environment, 89, 5-21. Meek, B., Loxton, D., Sparks, T., Pywell, R. Pickett, H. & Nowakowski, M. (2002). The effect of arable field margin composition on invertebrate diversity. Biological Conservation, 106, 259-271. Mitchley, J. (1994). Sward structure with regard to conservation. In: Haggar, R.J. and Peel, S. (Eds). Grassland Management and Nature Conservation, Occasional Symposium of the British Grassland Society, No. 28, pp. 43-53. Moonen, A.C. & Marshall, E.J.P. (2001) The influence of sown margin strips, management and boundary structure on herbaceous field margin vegetation in two neighbouring farms in southern England. Agriculture, Ecosystems & Environment, 86, 187-202. Økland, R.H. (1999). On the variation explained by ordination and constrained ordination axes. Journal of Vegetation Science, 10, 131-136. Økland, R.H. (2003). Partitioning the variation in a plot-by-species data matrix that is related to n sets of explanatory variables. Journal of Vegetation Science, 14, 693-700. Økland, R.H. & Eilertsen, O. (1994). Canonical Correspondence Analysis with variation partitioning: some comments and an application. Journal of Vegetation Science, 5, 117-126. Preston, C.D., Pearman, D.A. & Dines, T. (2002). New Atlas of the British Flora. Oxford University Press, Oxford. Pywell, R.F., Warman, E.A., Carvell, C., Sparks, T.H., Dicks, L.V., Bennett, D., Wright, A., Critchley, C.N.R. & Sherwood, A. (2005a). Providing foraging resources for bumblebees in intensively farmed landscapes. Biological Conservation, 121, 479-494. Pywell, R.F., Warman, E.A., Sparks, T.H., Nuttall, P., Hulmes, S., Hulmes, L., Wright, A., Saunders, P., Boyd, J., Taylor, A., Chapman, R., Critchley C.N.R. & Sherwood, A. (2005b). Effectiveness of new agri-environment schemes in providing foraging resources for bumblebees in intensively farmed landscapes. Report to Defra (this contract). CEH, Monks Wood. Rodwell, J.S. (ed.) (2000). British Plant Communities, Volume 5. Maritime Communities and Vegetation of Open Habitats. Cambridge University Press, Cambridge.

28 MA01017 CSS Grass Margins ADAS June 2005 Schippers, P. & Wouter, J. (2002). Modelling the effect of fertiliser, mowing, disturbance and width on the biodiversity of plant communities of field boundaries. Agriculture, Ecosystems & Environment, 93, 351-365. Smart, S.M., Firbank, L.G., Bunce, R.G.H. & Watkins, J.W. (2000). Quantifying changes in abundance of food plants for butterfly larvae and farmland birds. Journal of Applied Ecology, 37, 398-414. Smith, H. & MacDonald, D.W. (1989). Secondary succession on extended arable fields: its manipulation for wildlife benefit and weed control. Brighton Crop Protection Conference - Weeds 1989, pp. 1063-1068. British Crop Protection Council, Farnham. Smith, H., Feber, R. & Macdonald, D. (1999a) Sown field margins: why stop at grass? Field margins and buffer zones: Ecology, Management and Policy, (eds N. D. Boatman, D. H. K. Davies, K. Chaney, R. Feber, G. R. de Snoo, and T. H. Sparks) Aspects of Applied Biology 54, 275-282. Smith, H., Firbank, L.G. & Macdonald, D.W. (1999b). Uncropped edges of arable fields managed for biodiversity do not increase weed occurrence in adjacent crops. Biological Conservation, 89, 107-111. Stace, C.A. (1997). New Flora of the British Isles, 2nd edition. Cambridge University Press, Cambridge. StatSoft, Inc. (2001). STATISTICA for Windows [Computer program manual]. Tulsa, Oklahoma. ter Braak, C.J.F. & Šmilauer, P. (1998). CANOCO reference manual and user’s guide to CANOCO for Windows: Software for canonical community ordination (version 4). Microcomputer Power, Ithaca, . Thomas, C.F.G. & Marshall, E.J.P. (1999). Arthropod abundance and diversity in differently vegetated margins of arable fields. Agriculture, Ecosystems & Environment, 72, 131- 144. Thomas, M.B., Wratten, S.D. & Sotherton, N.W. (1991). Creation of 'island' habitats in farmland to manipulate populations of beneficial arthropods: predator densities and emigration. Journal of Applied Ecology, 28, 906-917. van Dorp, D., Schippers, P. & van Groenendael, J.M. (1997). Migration rates of grassland plants along corridors in fragmented landscapes assessed with a cellular automation model. Landscape Ecology, 12, 39-50. West, T.M., Marshall, E.J.P., Westbury, D.B. & Arnold, G.M. (1999). Vegetation development on sown and unsown field boundary strips established in three Environmentally Sensitive Areas. Field margins and buffer zones: Ecology, Management and Policy, (eds N. D. Boatman, D. H. K. Davies, K. Chaney, R. Feber, G. R. de Snoo, and T. H. Sparks) Aspects of Applied Biology, 54, 257-262. Wilson, P.J. (1999). The Ecology and Identification of Rare Arable Plants Recorded in the Arable Stewardship Pilot Areas. MAFF, Starcross. Wilson, P.J. & King, M. (2003). Arable Plants – A Field Guide. English Nature / Wildguides Ltd. www.arableplants.fieldguide.co.uk.

29 MA01017 CSS Grass Margins ADAS June 2005

Appendices

Appendix 1. List of grass species from which a mixture should be sown in Option R3 (from Defra, 2003).

Agrostis capillaris Common Bent Alopecurus pratensis Meadow Foxtail Anthoxanthum odoratum Sweet Vernal-grass Cynosurus cristatus Crested Dog’s-tail Festuca ovina Sheep’s Fescue Festuca pratensis Meadow Fescue Bromus commutatus Meadow Brome Hordeum secalinum Meadow Barley Festuca arundinacea Tall Fescue Festuca rubra (subsp. commutata) Red Fescue Poa pratensis Smooth Meadow-grass Trisetum flavescens Yellow Oat-grass Phleum bertolonii Smaller Cat’s-tail Dactylis glomerata Cocksfoot Deschampsia flexuosa Wavy Hair-grass Poa trivialis Rough Meadow-grass Phleum pratense Timothy Briza media Quaking Grass

31 MA01017 CSS Grass Margins ADAS June 2005 Appendix 2. Target List of Rare Arable Species

NR = Nationally Rare, NS = Nationally Scarce, SCC = Species of Conservation Concern.

Scientific Name English Name Status Adonis annua Pheasant’s-eye NR Agrostemma githago Corncockle Ajuga chamaepitys Ground-pine NR Alyssum alyssoides Small Alison SCC Anagallis arvensis ssp. foemina Blue Pimpernel NS Anthemis arvensis Corn Chamomile Apera interrupta Dense Silky-bent NS Apera spica-venti Loose Silky-bent NS Briza minor Lesser Quaking-grass NS Bromus arvensis Field Brome Bromus secalinus Rye Brome Bunium bulbocastanum Greater Pignut NR Centaurea cyanus Cornflower NR Chrysanthemum segetum Corn Marigold Echium plantagineum Purple Viper's-bugloss NR Erodium moschatum Musk Stork's-bill NS Euphorbia exigua Dwarf Spurge Euphorbia platyphyllos Broad-leaved Spurge NS Filago lutescens Red-tipped Cudweed NR Filago pyramidata Broad-leaved Cudweed NR Fumaria densiflora Dense-flowered Fumitory NS Fumaria occidentalis Western Ramping-fumitory NR Fumaria parviflora Fine-leaved Fumitory NS Fumaria purpurea Purple Ramping-fumitory NS Fumaria vaillantii Few-flowered Fumitory NS Galeopsis angustifolia Red Hemp-nettle NS Galium tricornutum Corn Cleavers NR Gastridium ventricosum Nit-grass NS Hypochaeris glabra Smooth Cat's-ear NS Iberis amara Wild Candytuft NS Lathyrus aphaca Yellow Vetchling NS Lavatera cretica Smaller Tree-mallow NR Lithospermum arvense Field Gromwell SCC Lythrum hyssopifolium Grass-poly NR Misopates orontium Weasel’s-snout Myosurus minimus Mousetail Petroselinum segetum Corn Parsley Papaver argemone Prickly Poppy Papaver dubium ssp. lecoqii Babington’s Poppy Papaver hybridum Rough Poppy Polycarpon tetraphyllum Four-leaved Allseed NR Polygonum rurivagum Cornfield Knotgrass Ranunculus arvensis Corn Buttercup NS Ranunculus muricatus Prickly-fruited Buttercup NS Ranunculus parviflorus Small-flowered Buttercup Scandix pecten-veneris Shepherd's-needle NS 32 MA01017 CSS Grass Margins ADAS June 2005 Scientific Name English Name Status Silene gallica Small-flowered Catchfly NS Silene noctiflora Night-flowering Catchfly SCC Torilis arvensis Spreading Hedge-parsley NS Valerianella dentata Narrow-fruited Cornsalad SCC Valerianella rimosa Broad-fruited Cornsalad NR Veronica praecox Breckland Speedwell NR Veronica triphyllos Fingered Speedwell NR Veronica verna Spring Speedwell NR Vicia parviflora Slender Tare NS

33 MA01017 CSS Grass Margins ADAS June 2005 Appendix 3. Plant taxa classified as farmland bird, butterfly larva or bumblebee food plants. Bird and butterfly plants from Smart et al. (2000), bumblebee plants from observations during surveys of Arable Stewardship Pilot Scheme sites (Pywell et al., 2005a).

Taxon Farmland Bird Butterfly Larva Bumblebee

Agrostis capillaris 9 Alliaria petiolata 9 Anchusa arvensis 9 Anthoxanthum odoratum 9 Arctium agg. 9 Arctium lappa 9 Arctium minus 9 9 Arrhenatherum elatius 9 Artemisia vulgaris 9 Avena fatua 9 Avena sativa 9 Avena sp. 9 Ballota nigra 9 Barbarea vulgaris 9 Beta sp. 9 Beta vulgaris ssp. vulgaris 9 Borago officinalis 9 Brachypodium sylvaticum 9 Brassica napus 9 9 Brassica napus ssp. oleifera 9 9 Brassica nigra 9 Brassica oleracea 9 Brassica oleracea var. capitata 9 Brassica oleracea var. viridis 9 Brassica rapa 9 Brassica rapa ssp. rapa 9 Brassica sp. 9 Calystegia sepium 9 Capsella bursa-pastoris 9 9 Carduus nutans 9 Centaurea nigra 9 Centaurea scabiosa 9 Cerastium fontanum 9 Cerastium glomeratum 9 Chamerion angustifolium 9 Chenopodium album 9 Chenopodium ficifolium 9 Cirsium arvense 9 9 Cirsium palustre 9 9 Cirsium sp. 9 Cirsium vulgare 9 9 Convolvulus arvensis 9 Crataegus monogyna 9 Cynosurus cristatus 9 Dactylis glomerata 9 Deschampsia cespitosa 9 Dipsacus fullonum 9 Elytrigia repens 9 hirsutum 9 34 MA01017 CSS Grass Margins ADAS June 2005 Taxon Farmland Bird Butterfly Larva Bumblebee Festuca arundinacea 9 Festuca pratensis 9 9 Festuca rubra 9 Festuca sp. 9 Galeopsis tetrahit 9 Hedera helix 9 Helianthus annuus 9 9 Holcus lanatus 9 9 Holcus mollis 9 9 Hordeum vulgare/distichon 9 Kickxia spuria 9 Knautia arvensis 9* 9 purpureum 9 Lathyrus pratensis 9 Linaria vulgaris 9 Lolium multiflorum/perenne 9 9 Lotus corniculatus 9 9 Lotus pedunculatus 9 sylvestris 9 Medicago lupulina 9 9 Medicago sativa 9 Melilotus officinalis 9 Melilotus sp. 9 Onobrychis viciifolia 9 Papaver rhoeas 9 Persicaria hydropiper 9 Persicaria lapathifolia 9 9 Persicaria maculosa 9 9 Phacelia tanacetifolia 9 Picris echioides 9 Poa annua 9 9 Poa pratensis 9 Poa trivialis 9 Polygonum aviculare 9 Polygonum rurivagum 9 Potentilla anserina 9 Potentilla reptans 9 Prunus spinosa 9 Raphanus raphanistrum 9 Reseda lutea 9 9 Reseda luteola 9 Rosa sp. 9 Rubus fruticosus 9 9 Rumex acetosa 9 9 Rumex acetosella 9 9 Rumex conglomeratus 9 Rumex crispus 9 Rumex obtusifolius 9 Rumex sanguineus 9 Rumex sp. 9 Senecio erucifolius 9 Senecio jacobaea 9 9 Senecio vulgaris 9 9 Sinapis alba 9 Sinapis arvensis 9 Sisymbrium officinale 9 Sonchus arvensis 9 9 35 MA01017 CSS Grass Margins ADAS June 2005 Taxon Farmland Bird Butterfly Larva Bumblebee Sonchus asper 9 Sonchus oleraceus 9 Sonchus sp. 9 Spergula arvensis 9 Stachys sylvatica 9 Stellaria holostea 9 Stellaria media 9 Taraxacum officinale agg. 9 Trifolium arvense 9 Trifolium dubium 9 9 Trifolium pratense 9 9 9 Trifolium repens 9 9 9 Tripleurospermum inodorum 9 Triticum sp. 9 Urtica dioica 9 9 9 Vicia hirsute 9 Vicia sativa 9 Vicia sp. 9 Vicia tetrasperma 9 Viola arvensis 9 Zea mays 9

* additional species not listed in Smart et al. (2000).

36 MA01017 CSS Grass Margins ADAS June 2005 Appendix 4. Occurrence of taxa and other variables (% of sites) from quadrat records of controls, sown R3 pre-2002, R3 post-2001, R3 + F and naturally regenerated (NR) sites. * species sown in at least some sites. name English Name All sites Control R3 pre R3 post R3+F NR Bare ground 66.4 87.9 54.2 69.0 54.5 45.5 Litter 65.5 57.6 75.0 62.1 63.6 81.8 Seedlings 58.8 57.6 54.2 69.0 63.6 36.4 Bryophytes 52.1 51.5 54.2 41.4 68.2 45.5 5.0 0 4.2 6.9 9.1 9.1

Crops & crop volunteers Triticum sp. Wheat 40.3 78.8 20.8 37.9 18.2 18.2 Hordeum distichon Two-rowed Barley 14.3 39.4 0 10.3 0 9.1 Solanum tuberosum Potato 5.0 6.1 0 6.9 9.1 0 Avena sativa Oat 1.7 6.1 0 0 0 0 Brassica napus ssp. oleifera Oil-seed Rape 1.7 3.0 0 3.4 0 0 Brassica oleracea Cabbage 0.8 0 0 3.4 0 0 Vicia faba Broad Bean 0.8 0 0 3.4 0 0 Pisum sativum Garden Pea 0.8 0 0 3.4 0 0 Brassica rapa Turnip 0.8 0 0 0 0 9.1

Grasses & rushes Poa trivialis Rough Meadow-grass 66.4 48.5 *70.8 *72.4 *77.3 72.7 Lolium perenne Perennial Rye-grass 62.2 36.4 *58.3 *75.9 *77.3 81.8 Dactylis glomerata Cock's-foot 61.3 15.2 *100 *69.0 *77.3 63.6 Agrostis stolonifera Creeping Bent 52.1 15.2 *70.8 55.2 *72.7 72.7 Poa annua Annual Meadow-grass 51.3 57.6 29.2 75.9 50.0 18.2 Phleum pratense Timothy 47.9 6.1 *87.5 *65.5 *50.0 36.4 Festuca rubra Red Fescue 47.9 0 *75.0 *58.6 *95.5 9.1 Elytrigia repens Common Couch 43.7 42.4 58.3 41.4 31.8 45.5 Holcus lanatus Yorkshire-fog 42.9 12.1 *58.3 37.9 *59.1 81.8 Arrhenatherum elatius False Oat-grass 37.0 24.2 41.7 27.6 50.0 63.6 Anisantha sterilis Barren Brome 33.6 27.3 29.2 44.8 22.7 54.5 Cynosurus cristatus Crested Dog's-tail 33.6 0 *50.0 41.4 *68.2 9.1 Poa pratensis Smooth Meadow-grass 30.3 6.1 *33.3 *41.4 50.0 27.3 Festuca pratensis Meadow Fescue 30.3 0 *37.5 *48.3 *54.5 9.1 Bromus hordeaceus Soft-brome 26.9 18.2 29.2 24.1 27.3 54.5 Agrostis capillaris Common Bent 25.2 0 *25.0 *27.6 *54.5 36.4 Holcus mollis Creeping Soft-grass 16.8 15.2 12.5 20.7 13.6 27.3 Alopecurus myosuroides Black-grass 11.8 6.1 12.5 20.7 9.1 9.1 Alopecurus pratensis Meadow Foxtail 11.8 3.0 *8.3 *13.8 *18.2 27.3 Festuca arundinacea Tall Fescue 10.9 0 *25.0 *13.8 *13.6 0 Avena fatua Wild-oat 10.1 18.2 0 17.2 4.5 0 Phleum bertolonii Smaller Cat's-tail 8.4 0 12.5 *13.8 *9.1 9.1 Anthoxanthum odoratum Sweet Vernal-grass 6.7 0 0 *3.4 *27.3 9.1 Lolium multiflorum Italian Rye-grass 5.9 9.1 4.2 6.9 4.5 0 Agrostis gigantea Black Bent 5.9 6.1 8.3 6.9 0 9.1 Festuca ovina Sheep's-fescue 5.0 0 *4.2 *6.9 *13.6 0 Trisetum flavescens Yellow Oat-grass 5.0 0 0 *6.9 *18.2 0 Bromus sp. Brome 4.2 6.1 0 0 *13.6 0 Alopecurus geniculatus Marsh Foxtail 4.2 0 4.2 10.3 0 9.1 x Festulolium loliaceum Hybrid Fescue 3.4 6.1 4.2 3.4 0 0 Agrostis sp. Bent-grass 3.4 0 *12.5 3.4 0 0

37 MA01017 CSS Grass Margins ADAS June 2005 Latin name English Name All sites Control R3 pre R3 post R3+F NR Deschampsia cespitosa Tufted Hair-grass 3.4 0 0 0 4.5 27.3 Bromus commutatus Meadow Brome 2.5 3.0 4.2 0 4.5 0 Juncus bufonius Toad Rush 2.5 3.0 0 3.4 4.5 0 Poa compressa Flattened Meadow-grass 2.5 0 *8.3 3.4 0 0 Vulpia bromoides Squirreltail Fescue 2.5 0 0 6.9 4.5 0 Phleum sp. Cat's-tail 2.5 0 0 *3.4 4.5 9.1 Bromus racemosus Smooth-brome 1.7 6.1 0 0 0 0 Bromus lepidus Slender Soft-brome 1.7 0 4.2 3.4 0 0 Juncus effusus Soft-rush 1.7 0 4.2 0 0 9.1 Hordeum murinum Wall Barley 1.7 0 0 3.4 4.5 0 Avena sp.Oat 1.7 0 0 3.4 0 9.1 Phalaris canariensis Canary-grass 1.7 0 4.2 3.4 0 0 Anisantha diandra Great Brome 0.8 3.0 0 0 0 0 Apera spica-venti Loose Silky-bent 0.8 3.0 0 0 0 0 Bromopsis ramosa Hairy-brome 0.8 3.0 0 0 0 0 Elymus caninus Bearded Couch 0.8 3.0 0 0 0 0 Brachypodium sylvaticum False Brome 0.8 0 4.2 0 0 0 Phragmites australis Common Reed 0.8 0 4.2 0 0 0 Agrostis canina Velvet Bent 0.8 0 0 0 *4.5 0 Poa sp. Meadow-grass 0.8 0 *4.2 0 0 0 Eleocharis sp. Spike-rush 0.8 0 0 3.4 0 0 Poaceae sp. Unidentified Grass 0.8 0 0 3.4 0 0 Helictotrichon pubescens Downy Oat-grass 0.8 0 0 0 *4.5 0 Hordeum secalinum Meadow Barley 0.8 0 0 0 4.5 0 Phalaris arundinacea Reed Canary-grass 0.8 0 0 0 4.5 0 Brachypodium pinnatum Tor-grass 0.8 0 0 0 0 9.1 Juncus conglomeratus Compact Rush 0.8 0 0 0 0 9.1

Forbs Galium aparine Cleavers 59.7 54.5 75.0 55.2 45.5 81.8 Cirsium arvense Creeping Thistle 55.5 27.3 70.8 55.2 63.6 90.9 Urtica dioica Common Nettle 43.7 27.3 45.8 41.4 50.0 81.8 Ranunculus repens Creeping Buttercup 38.7 6.1 62.5 20.7 *63.6 81.8 Trifolium repens White Clover 34.5 3.0 50.0* 41.4 *54.5 36.4 Cirsium vulgare Spear Thistle 31.1 9.1 25.0 48.3 40.9 45.5 Polygonum aviculare Knotgrass 28.6 39.4 16.7 37.9 27.3 0 Rumex obtusifolius Broad-leaved Dock 28.6 3.0 62.5 27.6 31.8 27.3 Taraxacum officinale Dandelion 26.9 15.2 25.0 27.6 40.9 36.4 Heracleum sphondylium Hogweed 26.9 9.1 41.7 24.1 *22.7 63.6 Veronica arvensis Wall Speedwell 24.4 18.2 29.2 31.0 22.7 18.2 Stellaria media Common Chickweed 22.7 18.2 20.8 41.4 13.6 9.1 Sonchus asper Prickly Sow-thistle 22.7 3.0 16.7 48.3 22.7 27.3 Geranium dissectum Cut-leaved Crane's-bill 21.0 15.2 20.8 24.1 27.3 18.2 Anthriscus sylvestris Cow Parsley 17.6 18.2 25.0 17.2 9.1 18.2 Tripleurospermum inodorum Scentless Mayweed 16.8 12.1 8.3 37.9 13.6 0 Plantago major Greater Plantain 16.8 0 25.0 27.6 18.2 18.2 Capsella bursa-pastoris Shepherd's-purse 15.1 6.1 0 48.3 9.1 0 Achillea millefolium Yarrow 15.1 0 12.5 6.9 *54.5 9.1 Convolvulus arvensis Field Bindweed 14.3 15.2 12.5 17.2 9.1 18.2 Matricaria discoidea Pineappleweed 14.3 9.1 16.7 27.6 4.5 9.1 Viola arvensis Field Pansy 13.4 15.2 8.3 17.2 18.2 0 Veronica persica Common Field-speedwell 12.6 15.2 0 31.0 4.5 0 Lapsana communis Nipplewort 12.6 15.2 0 20.7 13.6 9.1 Cerastium fontanum Common Mouse-ear 12.6 0 8.3 13.8 18.2 45.5 Prunella vulgaris Selfheal 12.6 0 8.3 3.4 *50.0 9.1

38 MA01017 CSS Grass Margins ADAS June 2005 Latin name English Name All sites Control R3 pre R3 post R3+F NR Lotus corniculatus Common Bird's-foot-trefoil 12.6 0 4.2 13.8 *40.9 9.1 Fallopia convolvulus Black-bindweed 11.8 18.2 4.2 20.7 4.5 0 Chenopodium album Fat-hen 11.8 3.0 8.3 27.6 13.6 0 Centaurea nigra Common Knapweed 11.8 0 0 0 *63.6 0 Persicaria maculosa Redshank 10.9 18.2 0 17.2 9.1 0 Sisymbrium officinale Hedge Mustard 10.9 6.1 0 20.7 18.2 9.1 Sonchus oleraceus Smooth Sow-thistle 10.1 3.0 4.2 13.8 18.2 18.2 Trifolium pratense Red Clover 10.1 0 4.2 10.3 *36.4 0 Leucanthemum vulgare Oxeye Daisy 10.1 0 4.2 3.4 *45.5 0 Artemisia vulgaris Mugwort 9.2 12.1 4.2 6.9 18.2 0 Epilobium sp. Willowherb 9.2 6.1 0 6.9 18.2 27.3 Anagallis arvensis ssp. arvensis Scarlet Pimpernel 9.2 3.0 4.2 17.2 9.1 18.2 Plantago lanceolata Ribwort Plantain 9.2 0 4.2 6.9 *36.4 0 Matricaria recutita Scented Mayweed 8.4 6.1 4.2 20.7 4.5 0 Glechoma hederacea Ground-ivy 8.4 3.0 20.8 10.3 0 9.1 Picris echioides Bristly Oxtongue 8.4 3.0 8.3 13.8 9.1 9.1 Ranunculus acris Meadow Buttercup 8.4 0 4.2 3.4 *27.3 18.2 Senecio vulgaris Groundsel 8.4 0 0 24.1 13.6 0 Geranium molle Dove's-foot Crane's-bill 7.6 6.1 8.3 3.4 9.1 18.2 Myosotis arvensis Field Forget-me-not 7.6 3.0 8.3 13.8 4.5 9.1 Rumex crispus Curled Dock 7.6 0 16.7 3.4 13.6 9.1 Veronica serpyllifolia Thyme-leaved Speedwell 6.7 6.1 0 6.9 4.5 27.3 Epilobium hirsutum Great Willowherb 6.7 3.0 12.5 0 13.6 9.1 Hedera helix Ivy 6.7 0 4.2 6.9 13.6 18.2 Daucus carota Wild Carrot 5.9 3.0 4.2 3.4 *18.2 0 Trifolium dubium Lesser Trefoil 5.9 0 16.7 0 13.6 0 Senecio jacobaea Common Ragwort 5.9 0 8.3 3.4 9.1 18.2 Aethusa cynapium Fool's Parsley 5.0 12.1 0 6.9 0 0 Arctium minus Lesser Burdock 5.0 3.0 4.2 6.9 4.5 9.1 Papaver rhoeas Common Poppy 5.0 3.0 0 13.8 4.5 0 Galium verum Lady's Bedstraw 5.0 0 0 3.4 *22.7 0 Rumex acetosa Common Sorrel 5.0 0 0 0 *22.7 9.1 Calystegia sepium Hedge Bindweed 4.2 6.1 8.3 0 0 9.1 Lamium purpureum Red Dead-nettle 4.2 6.1 0 6.9 4.5 0 Galium mollugo Hedge Bedstraw 4.2 3.0 4.2 3.4 9.1 0 Vicia cracca Tufted Vetch 4.2 3.0 4.2 3.4 *9.1 0 Silene latifolia White Campion 4.2 3.0 4.2 0 13.6 0 Epilobium ciliatum American Willowherb 4.2 0.0 8.3 6.9 4.5 0 Sinapis arvensis Charlock 3.4 6.1 0 6.9 0 0 Geranium robertianum Herb-robert 3.4 3.0 8.3 3.4 0 0 Atriplex patula Common Orache 3.4 3.0 0 10.3 0 0 Ballota nigra Black Horehound 3.4 3.0 0 3.4 9.1 0 Chaerophyllum temulum Rough Chervil 3.4 3.0 0 3.4 *9.1 0 Rumex sanguineus Wood Dock 3.4 0 12.5 0 0 9.1 Rumex sp. Dock 3.4 0 4.2 6.9 4.5 0 Trifolium hybridum Alsike Clover 3.4 0 4.2 6.9 *4.5 0 Medicago lupulina Black Medick 3.4 0 4.2 0 *13.6 0 Spergula arvensis Corn Spurrey 3.4 0 0 6.9 9.1 0 Epilobium parviflorum Hoary Willowherb 3.4 0 0 6.9 4.5 9.1 Aphanes arvensis Parsley-piert 2.5 3.0 4.2 0 4.5 0 Epilobium montanum Broad-leaved Willowherb 2.5 3.0 0 3.4 0 9.1 Alliaria petiolata Garlic Mustard 2.5 0 12.5 0 0 0 Descurainia sophia Flixweed 2.5 0 4.2 6.9 0 0 Pulicaria dysenterica Common Fleabane 2.5 0 4.2 0 4.5 9.1 Vicia tetrasperma Smooth Tare 2.5 0 4.2 0 4.5 9.1

39 MA01017 CSS Grass Margins ADAS June 2005 Latin name English Name All sites Control R3 pre R3 post R3+F NR Persicaria lapathifolia Pale Persicaria 2.5 0 0 10.3 0 0 Coronopus squamatus Swine-cress 2.5 0 0 6.9 4.5 0 Knautia arvensis Field Scabious 2.5 0 0 0 *13.6 0 Kickxia spuria Round-leaved Fluellen 1.7 6.1 0 0 0 0 Aegopodium podagraria Ground-elder 1.7 3.0 0 3.4 0 0 Conium maculatum Hemlock 1.7 3.0 0 3.4 0 0 Tripleurospermum maritimum Sea Mayweed 1.7 3.0 0 3.4 0 0 Kickxia elatine Sharp-leaved Fluellen 1.7 3.0 0 0 4.5 0 Vicia hirsuta Hairy Tare 1.7 3.0 0 0 *4.5 0 Stachys sylvatica Hedge Woundwort 1.7 3.0 0 0 0 9.1 Odontites vernus Red Bartsia 1.7 0 4.2 3.4 0 0 Veronica chamaedrys Germander Speedwell 1.7 0 4.2 3.4 0 0 Lathyrus pratensis Meadow Vetchling 1.7 0 4.2 0 4.5 0 Salvia verbenaca Wild Clary 1.7 0 4.2 0 *4.5 0 Hypochaeris radicata Cat's-ear 1.7 0 4.2 0 0 9.1 Viola tricolor Wild Pansy 1.7 0 0 6.9 0 0 Crepis sp. Hawk's-beard 1.7 0 0 3.4 4.5 0 Pimpinella saxifraga Burnet-saxifrage 1.7 0 0 3.4 *4.5 0 Raphanus raphanistrum Wild Radish 1.7 0 0 3.4 4.5 0 Sonchus arvensis Perennial Sow-thistle 1.7 0 0 3.4 4.5 0 Vicia sativa Common Vetch 1.7 0 0 3.4 *4.5 0 Crepis capillaris Smooth Hawk's-beard 1.7 0 0 0 *9.1 0 Hieracium sp. Hawkweed 1.7 0 0 0 *9.1 0 Malva moschata Musk-mallow 1.7 0 0 0 *9.1 0 Plantago media Hoary Plantain 1.7 0 0 0 9.1 0 Ranunculus bulbosus Bulbous Buttercup 1.7 0 0 0 9.1 0 Rhinanthus minor Yellow-rattle 1.7 0 0 0 *9.1 0 Silene dioica Red Campion 1.7 0 0 0 *9.1 0 Epilobium tetragonum Square-stalked Willowherb 1.7 0 0 0 4.5 9.1 Papaver sp. Poppy 1.7 0 0 0 0 18.2 Vicia sepium Bush Vetch 1.7 0 0 0 0 18.2 Brassicaceae sp. Crucifer species 0.8 3.0 0 0 0 0 Clematis vitalba Traveller's-joy 0.8 3.0 0 0 0 0 Coronopus sp. Swine-cress 0.8 3.0 0 0 0 0 Fumaria officinalis Common Fumitory 0.8 3.0 0 0 0 0 Fumaria sp. Fumitory 0.8 3.0 0 0 0 0 Galium sp. Bedstraw 0.8 3.0 0 0 0 0 Geranium pusillum Small-flowered Crane's-bill 0.8 3.0 0 0 0 0 Gnaphalium sp. Cudweed 0.8 3.0 0 0 0 0 Mercurialis annua Annual Mercury 0.8 3.0 0 0 0 0 Myosotis discolor Changing Forget-me-not 0.8 3.0 0 0 0 0 Solanum nigrum Black Nightshade 0.8 3.0 0 0 0 0 Solanum sp. Nightshade 0.8 3.0 0 0 0 0 Carduus crispus Welted Thistle 0.8 0 4.2 0 0 0 Cerastium sp. Mouse-ear 0.8 0 4.2 0 0 0 Chamerion angustifolium Rosebay Willowherb 0.8 0 4.2 0 0 0 Leontodon autumnalis Autumn Hawkbit 0.8 0 4.2 0 0 0 Lotus pedunculatus Greater Bird's-foot-trefoil 0.8 0 4.2 0 0 0 Persicaria amphibia Amphibious Bistort 0.8 0 4.2 0 0 0 Persicaria bistorta Common Bistort 0.8 0 4.2 0 0 0 Cardamine sp. Bitter-cress 0.8 0 0 3.4 0 0 Crepis vesicaria Beaked Hawk's-beard 0.8 0 0 3.4 0 0 Dipsacus fullonum Wild Teasel 0.8 0 0 3.4 0 0 Euphorbia sp. Spurge 0.8 0 0 3.4 0 0 Gnaphalium uliginosum Marsh Cudweed 0.8 0 0 3.4 0 0

40 MA01017 CSS Grass Margins ADAS June 2005 Latin name English Name All sites Control R3 pre R3 post R3+F NR Iris sp.Iris 0.8 0 0 3.4 0 0 Lepidium draba Hoary Cress 0.8 0 0 3.4 0 0 Linaria sp. Toadflax 0.8 0 0 3.4 0 0 Rorippa sp. Nothern Yellow-cress 0.8 0 0 3.4 0 0 Sanguisorba minor Salad Burnet 0.8 0 0 3.4 0 0 Sonchus sp. Sow-thistle 0.8 0 0 3.4 0 0 Stellaria graminea Lesser Stitchwort 0.8 0 0 3.4 0 0 Veronica scutellata Marsh Speedwell 0.8 0 0 3.4 0 0 Veronica sp. Speedwell 0.8 0 0 3.4 0 0 Agrimonia eupatoria Agrimony 0.8 0 0 0 *4.5 0 Centaurea scabiosa Greater Knapweed 0.8 0 0 0 *4.5 0 Cerastium glomeratum Sticky Mouse-ear 0.8 0 0 0 4.5 0 Cichorium intybus Chicory 0.8 0 0 0 *4.5 0 Clinopodium vulgare Wild Basil 0.8 0 0 0 *4.5 0 Hypericum perforatum Perforate St. John's-wort 0.8 0 0 0 *4.5 0 glandulifera Indian Balsam 0.8 0 0 0 4.5 0 Iris foetidissima Stinking Iris 0.8 0 0 0 4.5 0 Lactuca virosa Great Lettuce 0.8 0 0 0 4.5 0 Leontodon hispidus Rough Hawkbit 0.8 0 0 0 *4.5 0 Malva sp. Mallow 0.8 0 0 0 *4.5 0 Malva sylvestris Common Mallow 0.8 0 0 0 4.5 0 Medicago sativa ssp. sativa Lucerne 0.8 0 0 0 4.5 0 Mentha arvensis Corn Mint 0.8 0 0 0 4.5 0 Pastinaca sativa Parsnip 0.8 0 0 0 *4.5 0 Persicaria sp. Knotgrass 0.8 0 0 0 4.5 0 Primula veris Cowslip 0.8 0 0 0 *4.5 0 Rumex acetosella Sheep's Sorrel 0.8 0 0 0 4.5 0 Scabiosa columbaria Small Scabious 0.8 0 0 0 *4.5 0 Senecio erucifolius Hoary Ragwort 0.8 0 0 0 4.5 0 Stellaria holostea Greater Stitchwort 0.8 0 0 0 4.5 0 Tragopogon pratensis Goat's-beard 0.8 0 0 0 *4.5 0 Trifolium campestre Hop Trefoil 0.8 0 0 0 4.5 0 Barbarea vulgaris Winter-cress 0.8 0 0 0 0 9.1 Bryonia dioica White Bryony 0.8 0 0 0 0 9.1 White Dead-nettle 0.8 0 0 0 0 9.1 Myosotis sp. Forget-me-not 0.8 0 0 0 0 9.1 Sagina procumbens Procumbent Pearlwort 0.8 0 0 0 0 9.1 Stellaria graminea Lesser Stitchwort 0.8 0 0 0 0 9.1

Trees & shrubs Rubus fruticosus Bramble 16.0 3.0 25.0 17.2 13.6 36.4 Fraxinus excelsior Ash 8.4 0 12.5 10.3 18.2 0 Prunus spinosa Blackthorn 6.7 6.1 12.5 6.9 4.5 0 Crataegus monogyna Hawthorn 6.7 0 12.5 10.3 0 18.2 Acer pseudoplatanus Sycamore 3.4 3.0 4.2 3.4 4.5 0 Quercus robur Pedunculate Oak 3.4 3.0 0 0 4.5 18.2 Quercus sp.Oak 2.5 3.0 4.2 0 0 9.1 Rubus sp. Bramble 1.7 0 8.3 0 0 0 Acer campestre Field Maple 1.7 0 4.2 0 4.5 0 Fagus sylvatica Beech 1.7 0 0 0 9.1 0 Carpinus betulus Hornbeam 1.7 0 0 0 4.5 9.1 Acer sp. Maples 0.8 3.0 0 0 0 0 Rosa canina Dog-rose 0.8 0 0 3.4 0 0 Salix sp. Willow 0.8 0 0 3.4 0 0 Alnus glutinosa Alder 0.8 0 0 0 4.5 0

41 MA01017 CSS Grass Margins ADAS June 2005 Latin name English Name All sites Control R3 pre R3 post R3+F NR Tilia x europaea Common Lime 0.8 0 0 0 4.5 0 Castanea sativa Sweet Chestnut 0.8 0 0 0 0 9.1 Salix caprea Goat Willow 0.8 0 0 0 0 9.1 Sambucus nigra Elder 0.8 0 0 0 0 9.1

Horsetails & ferns Equisetum arvense Field Horsetail 10.1 6.1 8.3 13.8 13.6 9.1 Equisetum sp. Horsetail 2.5 0 4.2 6.9 0 0 Pteridium aquilinum Bracken 1.7 0 8.3 0 0 0 Equisetum palustre Marsh Horsetail 0.8 0 0 0 4.5 0

42 MA01017 CSS Grass Margins ADAS June 2005 Appendix 5. Abbreviated species names used in ordination figures.

Abbreviation Latin name Abbreviation Latin name Achimill Achillea millefolium Persmacu Persicaria maculosa Agrocapi Agrostis capillaris Phlebert Phleum bertolonii Agrostol Agrostis stolonifera Phleprat Phleum pratensis Anisster Anisantha sterilis Phlesp. Phleum sp. Anthodor Anthoxanthum odoratum Picrechi Picris echioides Arrhelat Arrhenatherum elatius Pisusati Pisum sativum Brasoler Brassica oleracea Planlanc Plantago lanceolata Cardsp. Cardamine sp. Planmajo Plantago major Capsburs Capsella bursa-pastoris Planmedi Plantago media Centnigr Centaurea nigra Poaannu Poa annua Cerafont Cerastium fontanum Poaprat Poa pratensis Chenalbu Chenopodium album Poatriv Poa trivialis Cirsarve Cirsium arvense Polyavic Polygonum aviculare Corosqua Coronopus squamatus Prunvulg Prunella vulgaris Cynocris Cynosurus cristatus Pteraqui Pteridium aquilinum Dactglom Dactylis glomerata Raphraph Raphanus raphanistrum Dauccaro Daucus carota Ranrepe Ranunculus repens Desccesp Deschampsia cespitosa Ranuacri Ranunculus acris Dipsfull Dipsacus fullonum Rhinmino Rhinanthus minor Elytrepe Elytrigia repens Rubufrut Rubus fruticosus Fagusylv Fagus sylvatica Rumeacet Rumex acetosa Fallconv Fallopia convolvulus Rumeobtu Rumex obtusifolius Festarun Festuca arundinacea Senevulg Senecio vulgaris Festovin Festuca ovina Siledioi Silene dioica Festprat Festuca pratensis Silelati Silene latifolia Festrubr Festuca rubra Sisyoffc Sisymbrium officinale Galimoll Galium mollugo Soncaspe Sonchus asper Galiveru Galium verum Soncoler Sonchus oleraceus Geradiss Geranium dissectum Sperarve Spergula arvensis Heraspho Heracleum sphondylium Stelmedi Stellaria media Hiersp. Hieracium sp. Tilix eu Tilia x europaea Holclana Holcus lanatus Trifcamp Trifolium campestre Horddist Hordeum distichon Trifdubi Trifolium dubium Irisfoet Iris foetidissima Trifprat Trifolium pratense Lamipurp Lamium purpureum Trifrepe Trifolium repens Lapscomm Lapsana communis Tripinod Tripleurospermum inodorum Leucvulg Leucanthemum vulgare Trisflav Trisetum flavescens Lolipere Lolium perenne Tritsp. Triticum sp. Lotucorn Lotus corniculatus Veroarve Veronica arvensis Malvmosc Malva moschata Veropers Veronica persica Matrdisc Matricaria discoidea Veroserp Veronica serpyllifolia Medilupu Medicago lupulina Vicitetr Vicia tetrasperma Paparhoe Papaver rhoeas Violarve Viola arvensis

43 MA01017 CSS Grass Margins ADAS June 2005 Appendix 6. Summary data for rare species.

Species: Apera spica-venti No. of records: 1

Grid ref. Date Field margin type Soil texture NVCNo. of individs SK 67 91 9/07/2004 control SZL OV20/21 > 1000

Community data (means from three 4m2 quadrats): Total Dicotyledons Monocotyledons Annuals Perennials Cover 17.7* 1.2 1.0 1.3 1.0 Richness 4.3** 2.3 2.0 0.7 0.5

Soil properties: pH P (mg l-1) K (mg l-1)Mg (mg l-1) 6.4 34 138 134

* Includes crop (Triticum sp., 7.7% without crop). ** Does not include crop (5.3 otherwise).

44 MA01017 CSS Grass Margins ADAS June 2005