The Floral and Butterfly Diversity of Green Lanes

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BIOLOGICAL CONSERVATION

Biological Conservation 121 (2005) 579–584 www.elsevier.com/locate/biocon

Linear hotspots? The ﬂoral and butterﬂy diversity of green lanes

P.J. Croxton, J.P. Hann, J.N. Greatorex-Davies, T.H. Sparks *

NERC Centre for Ecology and Hydrology, Monks Wood, Abbots Ripton, Huntingdon, Cambridgeshire, PE28 2LS UK

Received 7 January 2004; received in revised form 4 June 2004; accepted 7 June 2004

Abstract

In this study we compared the vascular flora and butterfly fauna of green lanes, single hedged tracks and grass verges. Green lanes supported a significantly more diverse flora with >30% more species in a 200 m transect than other linear features. Indicator values confirmed the lower light and higher moisture levels expected in green lanes, but in this study did not confirm lower nitrogen levels or greater stress tolerator incidence. Significantly more butterfly species were recorded in the green lanes and butterfly abundance was more than double that along the other linear features. Greater numbers of butterfly species were associated with greater numbers of floral species and green lanes also supported more butterfly larval foodplants. Given the value of butterflies as indicator species reflecting both the quality of habitat in the countryside and changes in the environment these results combine to emphasise the biodiversity value of green lanes in addition to their historical and aesthetic value. Ó 2004 Elsevier Ltd. All rights reserved.

Keywords: Butterﬂies; Green lanes; Hedgerows; Linear features; Farmland

1. Introduction the diversity and abundance of wildlife which was evi- dent during the latter part of the 20th century (e.g. Rob- Agricultural modernisation in the post war period led inson and Sutherland, 2002). to a simplification of the rural landscape, especially in The value of the remaining hedgerows and other lin- parts of eastern England, as family run mixed farms ear features such as ditches, green lanes and tracks is converted and combined into large arable enterprises. especially important as they may be the only semi-natu- Increased mechanisation led to greater cropping effi- ral habitats left in many rural areas. Linear features ciency and, to facilitate this expansion, many miles of have been recognised as providing a wide range of func- hedgerows were grubbed out and the input of artificial tions for a variety of wildlife. They act as linkages be- fertilisers, pesticides and herbicides increased dramati- tween habitats (Burel, 1996), roost and shelter sites cally (Robinson and Sutherland, 2002). For example, (Hinsley and Bellamy, 2000), provide feeding resources between 1945 and 1994 over 50% of the hedgerows in (Croxton and Sparks, 2002) and increase the diversity Leighton Bromswold in Cambridgeshire, close to the of the areas in which they occur. In particular the rela- sites used in this study, were removed resulting in the tively complex structure of green lanes provide an envi- average field size increasing from 7.5 to 22 ha (Westma- ronment that is especially valued by bees (Croxton et al., cott and Worthington, 1997). This simplification of the 2002), and butterflies (Dover et al., 2000; Dover and landscape inevitably contributed to the reduction in Sparks, 2001). From a wildlife perspective the diversity of structure

* and habitats oﬀered by having a variety of linear fea- Corresponding author. Tel.: +44-1487-772-461; fax: +44-1487- 773-467. tures would oﬀer more niches and ensure the greatest E-mail address: [email protected] (T.H. Sparks). overall biodiversity. Green lanes may be among the

0006-3207/$ - see front matter Ó 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2004.06.008 580 P.J. Croxton et al. / Biological Conservation 121 (2005) 579–584 oldest linear features in a landscape and provide a rela- 2.2. Vegetation survey tively stable environment with a localised microclimate more akin to woodland (Dover et al., 1997). The flora The abundance of vascular plant species and bare within green lanes is also likely to be protected to a ground present along the three transects at each site greater extent from physical and chemical disturbance was recorded by one of us (JNGD) using the DAFOR than the flora adjacent to single hedgerows. (dominant, abundant, frequent, occasional, rare) scale Green lanes, despite having important biological, cul- and the mean species richness (number of species) for tural and aesthetic values, have no specific protection; each boundary type calculated. DAFOR values were although The Hedgerows Regulations 1997 (Anon, converted to a 0–5 score (0=absent, 1=rare, 5=domi- 1997) mention ‘‘a parallel hedge within 15 m of a hedge- nant) prior to analysis. All plant species were assigned row’’ as one of the criteria defining important hedge- Ellenberg values for British plants (Hill et al., 1999) rows. More specifically, green lanes have been for light (L), moisture (F), reaction (R), and nitrogen described as ‘‘unmetalled tracks between fields of varia- (N) and CSR values (Grime et al., 1988). Mean values ble width, which may or may not be sunk below or of these seven variables were then calculated for the raised above field level, and bounded on both sides by flora recorded on each transect at each site. The larval grass banks, hedgerows or dry stone walls’’ (Dover foodplants of butterflies were taken from Asher et al. et al., 2000) or ‘‘an unmetalled track which may or (2001). may not be a right of way for the public either on foot, horse, bicycle or motor vehicle, and which is usually 2.3. Butterfly survey bounded by hedges, walls or ditches’’ (Countryside Agency, 2000). For the purpose of this study only green Every transect at each site was visited 18 times be- lanes with continuous hedgerows along both sides were tween April and August and butterfly counts recorded considered. by one of us (JPH) using the Butterfly Monitoring To further understand the effect of the structure of Scheme methodology, i.e. all butterflies within a 5-m linear features on butterfly fauna and vascular flora, corridor along the transects were recorded (Pollard in this study we compared 10 green lanes in Cambridge- and Yates, 1993). Records of the small skipper Thymeli- shire with matched single hedges and grass verges. This cus sylvestris and large skipper Ochlodes venata were is at the heart of the intensive arable region in SE Brit- combined for statistical analysis due to uncertainty in ain, where woodland cover (ca. 2%) and other semi- the identification of some specimens on the wing. On natural features are sparse. Ellenberg indicator values some exceptionally wide lanes two parallel 5 m-wide (Hill et al., 1999) and Competitor-Stress Tolerator- counts were taken and averaged prior to analysis. Ruderal (CSR) indices (Grime et al., 1988) were also used to investigate differences between the three bound- 2.4. Statistical analysis ary types, and to examine if any differences in the vascular flora reflected environmental or plant strategy Correspondence analysis (CA) with downweighting differences. for rare species was used to determine the variation in both the plant and (log transformed) butterfly commu- nities between the transect types. Within the CA, inter- 2. Materials and methods site differences were eliminated using indicator variables as covariates (analogous to removing site effects in a uni- 2.1. Sites variate ANOVA). FriedmanÕs nonparametric two-way ANOVA test (sites and transect types as factors) was The study was carried out on 10 arable sites in Cam- used to analyse the vascular plant and butterfly data. bridgeshire in 1998 where the three types of linear features, double hedged green lanes, single hedges with a track or verge and grass tracks or verges without a 3. Results hedge (hereinafter referred to as verge), occurred in close proximity. All sites were separated from one another by 3.1. Vegetation at least 1 km. Standardisation of hedge height, track/ verge width and orientation was attempted, but inevita- A total of 194 plant species was recorded on the 30 bly choice was restricted and some verges associated transects. Mean species scores (for those species with a with single hedges were narrower than the green lane significant effect and/or with a mean >1.0), species rich- tracks. Verges were either bounded on both sides by ness, Ellenberg indices and CSR indices in each of the fields or by field/road combinations. At each site tran- three boundary types is summarised in Table 1. Statisti- sects of 200 m were measured on each of the three cally significant differences existed for 22 of the individ- boundary types and temporary markers installed. ual species. In all but four cases (Avena fatua, P.J. Croxton et al. / Biological Conservation 121 (2005) 579–584 581

Table 1 The mean scores (0–5) of individual species (showing signiﬁcance or with mean>1.0) and bare ground, mean species richness, mean Ellenberg and CSR indices for each transect type (n=10 of each) Green lane Single hedge Grass verge p Acer campestre Field Maple 1.0 0.6 0.1 Agrostis stolonifera Creeping Bent 2.5 1.8 1.5 Alopecurus pratensis Meadow Foxtail 0.8 0.4 0.0 * Anisantha sterilis Barren Brome 0.6 1.2 0.9 Anthriscus sylvestris Cow Parsley 2.2 1.6 1.5 Arrhenatherum elatius False Oat-grass 3.6 3.8 4.5 Avena fatua Wild Oat 0.0 0.0 0.3 * Brachypodium pinnatum Tor-grass 0.4 0.4 1.6 * Centaurea nigra Common Knapweed 1.6 0.6 1.5 Cirsium arvense Creeping Thistle 1.7 1.4 1.4 Cirsium vulgare Spear Thistle 1.0 0.5 0.3 * Convolvulus arvensis Field Bindweed 1.8 3.2 2.6 * Crataegus monogyna Hawthorn 2.7 3.2 0.3 ** Dactylis glomerata CockÕs-foot 3.0 2.0 2.9 Deschampsia cespitosa Tufted Hair-grass 0.9 0.1 0.2 ** Elytrigia repens Common Couch 2.7 2.5 2.7 Epilobium hirsutum Great Willowherb 1.0 0.6 0.6 Festuca arundinacea Tall Fescue 0.6 1.4 1.2 Festuca rubra Red Fescue 0.8 0.7 1.8 Fraxinus excelsior Ash 1.6 0.7 0.8 Galium aparine Cleavers 3.1 2.9 1.0 ** Geranium dissectum Cut-leaved CraneÕs-bill 1.0 0.6 1.3 Glechoma hederacea Ground-ivy 1.7 0.8 0.7 Heracleum sphondylium Hogweed 2.1 1.3 1.9 Holcus lanatus Yorkshire-fog 1.4 0.6 0.7 Hordeum secalinum Meadow Barley 1.0 0.0 0.4 Juncus inﬂexus Hard Rush 0.4 0.0 0.0 * Lolium perenne Perennial Rye-grass 3.8 2.7 3.4 Odontites vernus Red Bartsia 0.8 0.0 0.0 * Phleum pratense Timothy 1.4 0.2 0.5 * Picris echioides Bristly Oxtongue 1.6 0.7 1.4 Plantago lanceolata Ribwort Plantain 1.2 0.6 1.0 Plantago major Greater Plantain 2.4 1.2 1.4 * Poa pratensis Smooth Meadow-grass 0.7 0.0 0.0 * Poa trivialis Rough Meadow-grass 2.7 1.2 1.3 Potentilla anserina Silverweed 0.4 0.0 0.0 * Potentilla reptans Tormentil 0.3 0.6 0.8 Prunella vulgaris Selfheal 0.3 0.0 0.8 * Prunus domestica Plum 0.6 0.1 0.0 * Prunus spinosa Blackthorn 3.7 2.5 0.6 *** Ranunculus repens Creeping Buttercup 2.3 0.6 0.6 * Rhamnus carthartica Buckthorn 0.6 0.0 0.0 * Rosa sp. Dog Rose 2.1 2.0 0.8 Rubus fruticosus Blackberry 3.0 2.3 1.2 ** Rumex crispus Curled Dock 1.0 0.2 0.4 * Rumex sanguineus Wood Dock 1.6 0.7 0.9 Trifolium repens White Clover 0.8 0.0 0.0 * Urtica dioica Common Nettle 2.0 2.0 1.7 Bare ground 1.2 0.2 0.4

Species richness 48.1 32.3 35.9 ** Ellenberg L (light) 6.67 6.66 6.96 * Ellenberg F (moisture) 5.39 5.26 5.14 * Ellenberg R (reaction) 6.68 6.87 6.79 * Ellenberg N (nitrogen) 6.15 6.36 6.09 Competitor 0.48 0.51 0.47 Stress Tolerator 0.19 0.18 0.16 Ruderal 0.33 0.31 0.37 Species with English names in bold are larval foodplants of one or more of the butterﬂy species listed in Table 2. ‘‘p’’ refers to signiﬁcance from Friedman nonparametric two-way ANOVA. * p<0.05. ** p<0.01. *** p<0.001. 582 P.J. Croxton et al. / Biological Conservation 121 (2005) 579–584

1 of the within-site variation is explained, respectively, by the ﬁrst and second axes in Fig. 1). Each group of transect types shows some degree of spread along the x-axis but broadly separate into horizontal bands along the y-axis, indeed ANOVA of second axis scores reveals signiﬁcance between transect types (p<0.001). 0

CA axis 2 3.2. Butterﬂies

A total of 22 species of butterflies was recorded, and with the exception of one rare species were most numer- -1 ous in the green lanes (Table 2). Two species (meadow -1 0 1 brown Maniola jurtina and green-veined white Pieris CA axis1 napi) accounted for >50% of the butterfly abundance. Fig. 1. A CA of the vascular plant data on green lane (open circle), Statistical significance was achieved for nine species single hedge (double circle) and verge (solid circle) transects. and for species richness and abundance. Species richness was markedly greater in the green lanes than elsewhere Brachypodium pinnatum, Convolvulus arvensis and Pru- and abundance at least twice as great as the other tran- nella vulgaris) abundance was higher in the hedged transect types. Two species, the brown argus Aricia agestis sects, particularly the green lane. Species richness was and small copper Lycaena phlaeas, were only recorded significantly higher (at least 30% higher) in the green in green lanes. lanes. The verges were characterised by high light, low The CA ordination of the log transformed butterfly moisture and, surprisingly, low nitrogen indices. There abundance data (24% and 23% of the within-site varia- were no significant differences in CSR indices. tion in butterfly numbers is explained, respectively, on The variation in the vegetation was confirmed in the the first and second axes in Fig. 2) shows the broad sep- correspondence analysis ordination where separation of aration of the three transect types along the first axis. In- the three transect types is clearly shown (14% and 10% deed ANOVA on the scores of this axis confirms

Table 2 The mean abundance, species richness and overall abundance of butterﬂies for each transect type (n=10 of each) Green lane Single hedge Grass verge p Gonepteryx rhamni Brimstone 1.2 0.3 0.0 *** Aricia agestis Brown Argus 3.5 0.0 0.0 Polygonia c-album Comma 0.7 0.1 0.0 ** Polyommatus icarus Common Blue 3.9 0.2 0.1 Pieris napi Green-veined White 65.3 36.8 45.4 Pyronia tithonus Gatekeeper 22.3 9.6 8.0 Celastrina argiolus Holly Blue 1.4 0.5 0.5 Thymelicus sylvestris/Ochlodes venata Small Skipper/Large Skipper 17.9 4.4 10.5 ** Pieris brassicae Large White 21.5 12.5 6.0 *** Maniola jurtina Meadow Brown 68.4 27.8 20.6 Anthocharis cardamines Orange Tip 1.7 1.2 0.1 * Vanessa cardui Painted Lady 0.5 0.1 0.0 * Inachis io Peacock 13.6 4.6 3.0 ** Vanessa atalanta Red Admiral 2.2 1.0 0.5 * Aphantopus hyperantus Ringlet 9.6 4.4 2.9 Lycaena phlaeas Small Copper 0.3 0.0 0.0 Coenonympha pamphilus Small Heath 3.5 0.5 0.6 Aglais urticae Small Tortoiseshell 9.0 8.2 3.7 Pieris rapae Small White 12.4 3.0 3.3 Pararge aegeria Speckled Wood 2.2 0.2 0.0 ** Lasiommata megera Wall 0.0 0.2 0.0

Species richness 15.6 10.9 9.0 *** Abundance 260.6 115.6 105.2 * ‘‘p’’ refers to signiﬁcance from Friedman nonparametric two-way ANOVA. * p<0.05. ** p<0.01. *** p<0.001. P.J. Croxton et al. / Biological Conservation 121 (2005) 579–584 583

1.0 the greater abundance of larval foodplants, the more sheltered conditions and the greater structural variety within the green lane environment (Dover et al., 2000). The green lanes were the most variable of the boundary 0.5 types with considerable differences in the width of lane depending on their original use, the widest, for example, being old drove roads. Some of the green lanes had been CA axis 2 0.0 unmanaged for several years resulting in tall hedges forming a green tunnel, which restricted light levels and reduced the incidence of flowering ground flora and butterflies except specialists like the speckled wood -0.5 Pararge aegeria. -0.5 0.0 0.5 The species richness of the green lane flora, with a CA axis1 mean of 48 species, was circa 30% greater than that of Fig. 2. A CA of the butterfly data on green lane (open circle), single the grass verge vegetation (mean 36 species), and the sin- hedge (double circle) and verge (solid circle) transects. gle hedgerows (mean 32 species). The enhanced species richness may be anticipated by the varying conditions produced by the structure of green lanes. In addition significant differences between the transect types to the typical hedgerows species, narrow or unmanaged (p=0.02). green lanes may provide conditions for a suite of woodland species to occupy the inside of the lane and the 3.3. Butterfly-flora associations broad open green lanes provide habitat for grasses and forbs along the centre. Increased species richness is Species richness of butterflies was positively associ- likely to provide a greater variety of flower morphology ated with species richness of the vascular plants and extend the temporal availability of food resources to (r=0.58, p=0.001, n=30) and with the number of larval the benefit of both adult butterflies and their larval foodplants of the recorded butterflies (r=0.54, p=0.002, forms. Of the 17 most abundant dicotyledon species, n=30). In Table 1, larval foodplants of the recorded 14 were most abundant in the green lanes with one, Ur- butterflies are identified. Of these 19 species, 15 were tica dioica, being equally abundant in single hedges. Lar- most (or equally) numerous on the green lane sections. val foodplants were more common in the green lanes. ANOVA confirms a significantly (p=0.005) greater The green lanes and the single hedges both had a mean mean number of larval foodplants in green lanes (mean of 13 tree and shrub species suggesting that it is other 15.3) compared to single hedges (mean 9.4) or verges features such as ground flora, microclimate and struc- (11.4). In general, species were not restricted to locations ture influencing the butterfly fauna. where their larval foodplants occurred. This reflects the The double hedges of green lanes would be expected facts that: (i) foodplants may have occurred outside of to act as a buffer against pesticide, herbicide and ferti- the 200 m surveyed transect, (ii) that resources other liser drift passing to the inside of the green lanes. Peak than foodplants (e.g. nectar) are also important in distri- nitrogen concentration rates of 150 kghaÀ1 were re- bution and (iii) butterflies have lesser or greater degrees ported by Tsiouris and Marshall (1998) at the hedge/ of mobility. field interface but they point out that the buffering effect of hedges may offer some protection to the habitat be- yond and behind the hedges. This study found the Ellen- 4. Discussion berg nitrogen value was lower in the grass verge habitat (6.09) and the green lanes (6.15) than the single hedge- The results of this study clearly show that, of the rows (6.36). We had anticipated higher deposition on three linear features studied, the green lanes had the the verges, but the semi-permeable barrier effect of the greatest floral diversity, as expected from the results of hedge may have concentrated higher nitrogen deposition Croxton et al. (2002), and supported a greater diversity along the field side of hedges with small amounts pene- and abundance of butterflies, as suggested by Dover trating into green lanes. et al. (2000). An association between species richness Fields are frequently cultivated within 0.5 m or less of of plants and species richness of butterflies was found hedgerow boundaries and even this narrow band may in support of similar associations reported elsewhere receive herbicide treatment. In contrast, the interior of (e.g. Sparks and Parish, 1995). A number of reasons green lanes are generally cut somewhat infrequently, if may be put forward to account for the preference shown at all. Therefore, the structure of the vegetation is more by butterflies for the green lanes over the other linear complex thus providing more ovipositing opportunities, features, including the greater diversity of plant species, and enhancing the chance of survival of the eggs and 584 P.J. Croxton et al. / Biological Conservation 121 (2005) 579–584 larval stages of Lepidoptera and other invertebrates. Brown, K.S., Freitas, A.V.L., 2000. Atlantic forest butterflies: indica- The paucity of both plant and butterflies along the single tors for landscape conservation. Biotropica 32, 934–956. hedgerows emphasises that 6-m wide conservation strips Burel, F., 1996. Hedgerows and their role in agricultural landscapes. Critical Reviews in Plant Sciences 15 (2), 169–190. along field boundaries (e.g. Meek et al., 2002) may im- Countryside Agency, 2000. Out in the Country: Where You Can Go prove the diversity of these features. and What You Can Do. Booklet CA9. Countryside Agency The environmental conditions within green lanes Publication, Wetherby, West Yorkshire. are likely to be more stable than the conditions out- Croxton, P.J., Carvell, C., Mountford, J.O., Sparks, T.H., 2002. A side with reduced air movement between double comparison of green lanes and field margins as bumblebee habitat in an arable landscape. Biological Conservation 107, hedgerows and subsequently reduced evapotranspira- 365–374. tion leading to the higher moisture levels shown in Croxton, P.J., Sparks, T.H., 2002. A farm-scale evaluation of the Table 1. The height/width ratio of green lanes also influence of hedgerow cutting frequency on hawthorn (Crataegus play a role in affecting the diversity of butterflies uti- monogyna) berry yields. Agriculture, Ecosystems and Environment lising the habitat (Sparks et al., 1999). If the hedge- 93, 437–439. Dover, J.W., Sparks, T.H., 2001. Green lanes: biodiversity reservoirs rows in narrow green lanes are unmanaged they will in farmland? In: Barr, C., Petit, S. (Eds.), Hedgerows of the World: eventually form shady tunnels, suiting species such Their Ecological Functions in Different Landscapes. IALE (UK), as the speckled wood, but restricting the occurrence Lymm, pp. 241–250. of many butterfly species that thrive in more open Dover, J.W., Sparks, T.H., Clarke, S., Gobbett, K., Glossop, S., 2000. sunny habitats such as woodland rides and glades Linear features and butterflies: the importance of green lanes. Agriculture, Ecosystems and Environment 80, 227–242. (Greatorex-Davies et al., 1993). Dover, J.W., Sparks, T.H., Greatorex-Davies, J.N., 1997. The impor- The value placed on butterflies as indicator species to tance of shelter for butterflies in open landscapes. Journal of Insect reflect the quality of habitat in the countryside (Asher Conservation 1, 89–97. et al., 2001) and changes in the environment (Brown Greatorex-Davies, J.N., Sparks, T.H., Hall, M.L., Marrs, R.H., 1993. and Freitas, 2000) suggests that green lanes are a very The influence of shade on butterflies in rides of coniferised lowland woods in southern England and implications for conservation valuable landscape feature. Bumblebees also favour management. Biological Conservation 63, 31–41. the green lane habitat (Croxton et al., 2002) which, to- Grime, J.P., Hodgson, J.G., Hunt, R., 1988. Comparative Plant gether with the results reported here, emphasise the Ecology: A Functional Approach to Common British Species. importance of green lanes as linear hotspots at least Unwin Hyman, London. for nectar feeding invertebrates. Like all man-made fea- Hill, M.O., Mountford, J.O., Roy, D.B., Bunce, R.G.H., 1999. EllenbergÕs Indicator Values for British Plants. HMSO, Norwich. tures, green lanes need management to maximise their Hinsley, S.A., Bellamy, P.E., 2000. The influence of hedge structure, conservation value. A heterogenous approach to man- management and landscape context on the value of hedgerows to agement frequency of hedges and tracks would provide birds: a review. Journal of Environmental Management 60, 33– a range of floral, structural and microclimatic condi- 49. tions. Zero management is not a wise option since the Meek, B., Loxton, D., Sparks, T., Pywell, R., Pickett, H., Nowakowski, M., 2002. The effect of arable field margin lane would become swamped by rank grasses and even- composition on invertebrate biodiversity. Biological Conservation tually form a green tunnel. Some degree of disturbance 106, 259–271. to the track is recommended to encourage annuals and Pollard, E., Yates, T.J., 1993. Monitoring Butterflies for Ecology and increase the range of flowering plants. This study dem- Conservation. Chapman & Hall, London. onstrates that green lanes, in addition to their cultural Robinson, R.A., Sutherland, W.J., 2002. Post-war changes in arable farming and biodiversity in Great Britain. Journal of Applied and aesthetic value, can enhance local biodiversity with Ecology 39, 157–176. their suite of vascular plant species supporting butterfly Sparks, T.H., Hann, J.P., Greatorex-Davies, J.N., 1999. The influence species richness and abundance in an increasingly sim- of field boundary structure on butterflies. Aspects of Applied plified arable landscape. Biology 54, 235–240. Sparks, T.H., Parish, T., 1995. Factors affecting the abundance of butterflies in field boundaries in Swavesey Fens, Cambridgeshire, UK. Biological Conservation 73, 221–227. References Tsiouris, S., Marshall, E.P.J., 1998. Observations on patterns of granular fertiliser deposition beside hedges and its likely effects on Anon, 1997. The Hedgerow Regulations 1997. HMSO, London. the botanical composition of field margins. Annals of Applied Asher, J., Warren, M., Fox, R., Harding, P., Jeffcoate, G., Jeffcoate, S., Biology 132, 115–127. 2001. The Millenium Atlas of Butterflies in Britain and Ireland. Westmacott, R., Worthington, T., 1997. Agricultural Landscapes: A OUP, Oxford. Third Look. Countryside Commission, Cheltenham.