Species Action Plan THE SILVER-STUDDED BLUE Plebejus argus JUNE 1996 Compiled by: N.O.M.Ravenscroft and M.S.Warren Butterfly Conservation P.O.Box 444 Wareham Dorset BH20 5YA Tel: 01929 400209 This species action plan is an unpublished working document produced by Butterfly Conservation to focus and co-ordinate the conservation of the Silver-studded Blue in the UK over the next five to ten years. It has been prepared under the Action for Butterflies project which is funded by WWF-UK, English Nature, the Countryside Council for Wales and Scottish Natural Heritage. The Action Plan was prepared in consultation with the following organisations with the intention that they will participate in the actions outlined: English Nature, Countryside Council for Wales, Ministry of Agriculture, Fisheries and Food, Welsh Office Agriculture Department, Royal Society for the Protection of Birds, National Trust, the Wildlife Trusts (Suffolk, Sussex, Hampshire, Dorset, Devon and Cornwall), the Forestry Authority and the Forestry Commission (New Forest). Outline Paper (first draft): ....................................................................................... January 1996 Pathfinder Meeting: ..................................................................................................March 1996 Second Draft: ............................................................................................................March 1996 Final Draft: ...................................................................................................................June 1996 Acknowledgements We are grateful to the following for their comments at the Pathfinder meeting and / or on subsequent drafts; Andrew Nicholson, Rees Cox, Stuart Graham, Richard Cottle, Neil Hill (English Nature), Adrian Fowles (Countryside Council for Wales), Mike Clarke and Jon Gulliver (Forestry Commission, New Forest), Peter Goatham and Brian Pickess (Royal Society for the Protection of Birds), Nigel Bourn, Paul Kirkland and Peter Thornett (Butterfly Conservation), Robin Crane (The Wildlife Trusts), Jenny Joy, Chris Thomas (University of Leeds), Ken Wilmott, Simon Curson (Sussex Wildlife Trust), Clive Chatters (Hampshire Wildlife Trust), Liesje Birchenough (Dorset Wildlife Trust), Nigel Webb (Institute of Terrestrial Ecology), Ms A.E.Tarran (Ministry of Agriculture, Fisheries and Food) and Mr M.D.Giles (Welsh Office Agriculture Department). Butterfly Conservation Society (the British Butterfly Conservation Society) has an overriding objective to ensure a future for butterflies, moths and their habitats. In order to achieve this objective its aims are to raise public awareness of the plight of our butterflies and moths. to reduce the decline of commoner species and to increase the numbers of rarer species. to work with and advise other conservation groups, local bodies and agencies on techniques of land management which favour butterflies and moths and related wildlife. to acquire and manage habitats for butterflies and moths. to encourage the research (both at amateur and professional levels) on butterflies and moths. support and encourage butterfly and moth conservation world-wide. Registered Office of Butterfly Conservation: Shakespeare House, High Street, Dedham, Colchester, Essex, CO7 6DE. Registered in England No. 2206468 Registered Charity No. 254937. 2 CONTENTS Page Summary 4 Part 1 1.1 1.2 1.3 1.4 Overview Priority Statement Broad Objectives Legal Status Status and Level of Biological Knowledge Part 2 2.1 2.2 2.3 2.4 2.5 Biological Assessment Introduction Ecology Distribution and Population Limiting Factors Resume of Conservation to Date Part 3 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9 Actions and Work Programme Policy and Legislative Site Safeguard and Acquisition Land Management Species Protection and Licensing Advisory International Future Research, Survey and Monitoring Communications and Publicity Review Abbreviations 5 5 6 7 7 10 12 13 References Appendix 1 Appendix 2 5 15 15 15 16 16 16 17 17 18 19 Distribution in the UK Heathland management for the Silver-studded Blue 3 22 23 Summary The Silver-studded Blue Plebejus argus is a nationally scarce species that is included in the Guidelines for the Selection of Biological Sites of Special Scientific Interest (NCC 1989) and protected under Schedule 5 of the Wildlife and Countryside Act 1981 with respect to trade. Although it has declined substantially through much of its range, it is still locally abundant in heathlands in parts of southern England. A medium priority is afforded to the conservation action in this plan to protect and increase the number of Silver-studded Blue colonies in the UK. The British population has four subspecies: P. a. argus is the lowland heathland form found throughout its current range; P. a. caernensis occurs on calcicolous grassland in North Wales; P. a. cretaceus is now confined to calcicolous grassland in Dorset; and P. a. masseyi is a form found at a single moss site in Wales being extinct from Cumbria and Scotland. The genetic distinctiveness of these races is currently under investigation. The Silver-studded Blue has undergone a severe decline in range this century, estimated at 80%. It has become extinct in Scotland and northern England and throughout most of central, eastern and south-eastern England. It remains widespread only on the heaths of Dorset and Hampshire. It is regarded as an indicator of active lowland heathland management. The major threats to this species are heathland destruction, neglect or mis-management, combined with the fragmentation of its habitat which increases local extinction and prevents recolonisation. It requires foodplants growing alongside patches of open soil or in short turf and strong populations of one of two species of ants with which it has a relationship in its larval and pupal stages. These conditions are satisfied by habitats in early successional stages or those maintained by grazing. The major objectives of this plan are to ensure that existing populations are maintained through appropriate habitat management and to encourage restoration of habitat throughout its former range. The objectives of this plan will be achieved by encouraging beneficial land management on existing sites, especially those that occur on nature reserves; restoring and maintaining a network of habitat between existing sites and within its former range; and conducting strategic re-introductions. Further research and survey will be supported, particularly into the status and ecology of the species in relatively little described parts of its range. The Action Plan and the conservation of this butterfly will be publicised. Funds will be sought to ensure that proposed actions are implemented. The Action Plan covers the next ten years, will be monitored annually and reviewed in the year 2000 or such earlier time as the situation demands. 4 Part 1 Overview 1.1 PRIORITY STATEMENT The Silver-studded Blue is a nationally scarce species that is included in the Guidelines for the Selection of Biological Sites of Special Scientific Interest (NCC 1989). It is a butterfly of open ground supporting leguminous and ericaceous shrubs, and in Britain is primarily a heathland species (although it inhabits calcicolous grassland in small parts of its range). It has declined on heathland throughout Britain, becoming extinct in Scotland, absent in northern, central and south-east England and severely reduced in abundance elsewhere. Its main centres are the heaths of southern and south-west England, coastal Wales and parts of East Anglia. Remaining heathlands are highly fragmented and often too small to support colonies indefinitely as the species requires early successional, but transient habitats. Although the species has declined substantially in the last fifty years, it is still locally abundant in parts of its southern range. A medium priority should thus be afforded to implementing this National Plan, but a high priority may be given in those regions which have suffered a high rate of decline, or where it is reduced to a small number of populations. 1.2 BROAD OBJECTIVES 1. Maintain present range through suitable habitat management at a local and landscape scale. 2. Encourage restoration of suitable habitats in present and former range, concentrating on strategic sites and networks of habitats that will allow population spread and the maintenance of metapopulations. 1.3 LEGAL STATUS The Silver-studded Blue is listed on Schedule 5 of the 1981 Wildlife and Countryside Act for sale only. 5 1.4 Status and Level of Biological Knowledge Population Knowledge of -size Confined to southern parts of England, coastal Wales and parts of East Anglia. Most remaining colonies are small and isolated. About 50 colonies in N. Wales, 31 in Devon, 14 in Surrey, 10 in Suffolk and one in Shropshire, although most are small. Less detail for Dorset and Hampshire where it is more common, and Sussex, Norfolk and Cornwall. Few extensive tracts of habitat remain. -trend, numbers Recorded decline is severe, based on knowledge of 10km squares. Decline in numbers is continuing in some areas. Smaller populations are most vulnerable to extinction. -trend, range Severe contraction in range this century (65%). Now extinct throughout Scotland, northern England and most of central, eastern and south-eastern England. -status Good in most areas. -trends Good. -conservation requirements Good. Responds well to correct management. 6 Part 2 Biological Assessment 2.1 INTRODUCTION The Silver-studded Blue Plebejus argus has undergone a severe contraction in range throughout Britain and reductions in abundance in its remaining distribution (Heath et al 1984, Thomas 1993). It occurs in a variety of habitats: lowland acid heathland (dry and wet), calcicolous grassland, sand dunes and lowland mosses (Emmet & Heath 1990). Heathland is the most commonly used habitat in Britain and supports most of the population; colonies on this habitat are the nominate subspecies P. argus argus. Populations on calcicolous grasslands are now restricted to two locations: Great Orme's Head in North Wales (where P. argus caernensis occurs) and the Isle of Portland in Dorset (P. argus cretaceus). Colonies on the limestone of the Dulas valley in North Wales resulted from introductions made 50 years ago (Marchant 1956). One mossland population remains (in North Wales) although the species was once widespread in this habitat, especially in Cumbria (P. argus masseyi). Extinct populations in Scotland are thought to have resembled P. a. masseyi (Thomson 1980). The classification of these subspecies is based primarily on the size and colouration of adults (Emmet & Heath 1990), but their status is uncertain (Dennis 1977, Thomas 1985a). The genetics of isolated populations and subspecies are currently being studied. 2.2 ECOLOGY The Silver-studded Blue occurs at high densities in closed colonies (Thomas 1985b, Ravenscroft 1990a, Emmet & Heath 1990, Lewis et al. 1995). The butterfly is univoltine in Britain, flying from July to August in most years, although in some years it may emerge in mid-June (Heath, Pollard & Thomas 1984). Calcicolous populations usually emerge in June in N. Wales (Thomas 1985a). In parts of central and southern Europe it is bivoltine, emerging as early as May (Higgins & Riley 1970). The flight period lasts 4-5 weeks and numbers reach pronounced peaks within 2-3 weeks of emergence at most sites studied. The species exhibits protrandry, with males emerging before females. Both sexes are highly sedentary and most individuals move less than 50m over their lifespan (Thomas 1985b, Ravenscroft 1990a, Lewis et al. 1995). Adults take nectar from a variety of plants depending on their availability (Bell Heather Erica cinerea, Cross-leaved Heath Erica tetralix, Bramble Rubus fruticosus agg. and Bird's-foot Trefoil Lotus corniculatus) but are limited to the flowers of heathers at most heathland sites (Ravenscroft 1990a, Joy 1992, Wilmott 1994). Females feed more than males. Adults may congregate in conspicuous roosts, often on heather bushes or grass tussocks, where they roost head downwards overnight. Females lay eggs singly close to the ground on a variety of plants, but usually on the woody stems of heathers (Ling Calluna vulgaris, Bell Heather and Cross-leaved Heath) or Gorses (Ulex spp.)on heathlands (Thomas 1985a, Ravenscroft 1990a, Joy 1992, Wilmott 1994) and, especially in calcicolous habitats, on moss, lichens, twigs and leaf litter beneath prospective foodplants (Thomas 1985a). The egg is the overwintering stage. They are usually laid around the edges of short vegetation bordering patches of bare ground (Thomas 1985a). Eggs are usually laid <7cm above the ground on heathlands in N. Wales (Thomas 1985a), <7cm above the ground on heathland in Shropshire (Joy 1992), <10cm above the ground on 7 heathland in Suffolk (Ravenscroft 1986) and <3cm above the ground on calcicolous grassland in N. Wales (Thomas 1985a). Exceptions have been noted at some sites, and females may sometimes lay eggs high on grasses or in clusters under the fronds of sparse Bracken Pteridium aquilinum at heights in excess of 30cm (Ravenscroft 1986, Mendel & Parsons 1987). No detailed work has been done in the Dorset and Hampshire strongholds but the egg laying requirements are considered to be slightly less exacting in these southern locations. Larvae hatch in Spring and are able to feed on a wide range of plants from three families (the Leguminosae, Ericaceae and Cistaceae) (Thomas 1993). On heathland and moss sites the important foodplants appear to be Bell Heather, Cross-leaved Heath, Gorse, Ling, and probably Bird's-foot Trefoil where this occurs on grassy sites (heathland larvae prefer this last plant in captivity, Thomas 1985a) (Thomas 1985a,b, Joy 1992, Wilmott 1994). On calcicolous grassland, Bird's-foot Trefoil, Rock Rose (Helianthemum nummularium) and Hoary Rock-rose (H. canum) are used in North Wales (and very rarely Thyme, Thymus drucei) (Thomas 1985a) and Bird’s-foot Trefoil, Rock-rose and Horseshoe Vetch (Hippocrepis comosa) in Dorset (Warren 1986, Wilmott 1994). On Continental Europe, a wider range of plants may be used, including Bilberry (Vaccinium myrtillus), Broom (Cytisus scoparius) (also used in the U.K., e.g. at Prees Heath, C.D. Thomas pers. comm.) and Halimium halimifolium (Higgins & Riley 1970, Jordano et al 1992). In all parts of the range, larvae feed primarily on the buds, flowers, growing tips or young leaves of plants. In the UK, larvae are fully developed (about 13mm in length) by June. Throughout development, the Silver-studded Blue has a highly developed relationship with ants which is maintained by the production of sugar-rich liquid by the larvae and pupa. Females respond to the presence of ants when selecting sites to lay eggs (Jordano et al 1992). This may explain the tendency of females to lay on Bracken at some sites as this plant has extra-floral nectaries attended by ants (Mendel & Parsons 1987). Larvae have a pair of prominent reversible tentacular organs on the abdomen which attract and excite ants (perhaps by emitting chemicals) and small glands (pore cupola organs) scattered over the body, some of which may secrete sugar. They also have a nectary organ (Newcomer's organ) on the abdomen which develops in the third instar and secretes sugar and amino acids (Malicky 1969, Fiedler 1989). Ants gather droplets secreted from the pre-cupola and Newcomer's organs. Larvae are also covered with droplets when they hatch, pupae exude droplets and adults emerge covered in droplets (Thomas 1985a, Ravenscroft 1990a, Jordano & Thomas 1992). Ants tend larvae from hatching and through pupation. Full-grown larvae can be found in the field resting on top of ant nests, in ant nests or under stones used by ants (Thomas 1985b, Ravenscroft 1990a, Jordano et al 1992). On calcicolous sites in Dorset, larvae are thought to spend most of their time in ant nests and emerge to feed by night (Wilmott 1994). The ecology of the early instars is less clear. In captivity, ants are attracted to hatching larvae, which may be carried off after emerging (Ravenscroft 1990a, Jordano & Thomas 1992) and first and second instar have been found in ant chambers in the field (Wilmott 1994). It seems likely that most larvae are taken into ant nests where they rest, emerging only to feed. Others are tended where they occur on foodplants (Jordano et al 1992). Pupae are tended by ants under stones, under mats of prostrate vegetation such as rabbit-cropped gorse, small holes in the ground, in earthen cells constructed by ants or in ant nests (Thomas 1993, Jordano et al 1992, Emmet & Heath 1990). Although pre-pupal larvae are not thought to seek out ants 8 (Jordano et al 1992), a small nest may be established around the pupa (Emmet & Heath 1990). Adults emerge covered in ants which remain to gather droplets from their surfaces. The benefit of this relationship to the Silver-studded Blue is probably protection from parasites and predators, as has been demonstrated with closely related lycaenids (Pierce et al. 1987). Ants gain sugar and nutrition in return and there is evidence that ants which attend such larvae have higher survival rates (Fiedler & Saam 1995). The ants associated with the Silver-studded Blue are the common black ants Lasius niger and L. alienus and the relationship appears to be Lasius-specific, although ants of other genera will tend later instars in captivity (Jordano & Thomas 1992). In N. Wales, L. alienus is the main ant on limestone and L. niger on heathlands (Thomas 1985a). On Suffolk heathlands, the species used is site specific; L. niger is used at most sites but L. alienus takes over where L. niger is absent on the driest heaths (Ravenscroft 1986). Lasius flavus (the Yellow Meadow Ant) responds to larvae in captivity (Ravenscroft 1990a, Jordano & Thomas 1992) and is abundant on some sites (Ravenscroft 1986), but has not been recorded with larvae in the field. Habitat requirements The Silver-studded Blue requires the presence of Lasius sp. ants, open ground for breeding and either bare soil or short vegetation in all habitats in Britain. Choice of habitat varies from region to region and this may reflect differences in ant densities. Colonies in southern England may survive in places where heathers have become leggy, exposing bare ground beneath (Thomas & Webb 1984). On heathlands and calcicolous grassland, adult densities are correlated positively with a variety of indicators of the early stages of heathland and negatively with vegetation height, although the overall relationship appears to be with ant density (Jordano et al 1992, Thomas 1985b, Ravenscroft 1990a). The preferred conditions produce warmer microclimates at ground level for larvae; a factor that is especially important towards the north of the species' range (Thomas 1993). (At the southern limit in Spain taller vegetation is preferred which reduces ground temperatures, Jordano et al 1992). A warm microclimate is probably even more important to maintain the nest-building and foraging activities of Lasius ants. Densities of adults are closely correlated with those of ant nests (Ravenscroft 1990a) as is the distribution of eggs, larvae and pupae (Jordano et al (1992). The highest densities of butterflies and ants in Suffolk occur where Bell Heather is most abundant (6-30% ground cover) or where it replaces Ling altogether (Ravenscroft 1986). Butterflies do not occur where ant nest density falls below 30 nests/ha and ants are largely absent (<10 nests/ha) from dense or mature Ling dominated heathlands (>80% cover) (Ravenscroft 1986, 1992a). In West Sussex, the butterfly occurs almost exclusively where Bell Heather is abundant (Crane 1995). In Surrey, the butterfly is found in the presence of Erica heather and 5-15% bare ground (Wilmott 1994). Bell heather therefore appears to represent a good indicator of suitable conditions in many areas. In Dorset, however, the Silver-studded Blue is most often associated with humid and wet heath where Cross-leaved Heath is dominant and Bell Heather absent, but homogenous stands of Ling remain unsuitable. On calcicolous grassland in N. Wales, the Silver-studded Blue is absent from sites where ants are also largely absent (Jordano et al 1992). Early successional stages, and those held in check by grazing (nowadays mainly by rabbits in most areas, although the New Forest is pony/cattle grazed), are preferred in all habitats. On 9 heathland, there is a range of suitable conditions, from nutrient-deficient sandy soils to peaty soils. Most heathland colonies exist on sites that have been either recently disturbed, such as sand-pits, quarries and firebreaks, or burnt (Thomas 1985b, Ravenscroft 1990a, Wilmott 1994). Conditions usually become suitable after 2-5 years of re-growth. In the absence of traditional management, such as cutting of heathers and invasive scrub, grazing of domestic animals, or burning to encourage young growth for livestock, suitable conditions on heathlands are short-lived, perhaps 5-10 years (Thomas 1985b). Exceptions occur where heathland is grazed by rabbits, which maintain open, short vegetation; in wet hollows on humid heaths where vegetation succession is slowed by the prevailing conditions; and also in extreme southern counties like Dorset and Hampshire where slightly taller vegetation may be tolerated. On calcicolous grassland, conditions persist for much longer, and on Great Orme's Head continuous sheep and goat grazing may maintain suitable conditions indefinitely, although the butterfly may be excluded from areas that are too heavily grazed (Thomas 1985b). Ideal conditions are found in 2-7cm tall vegetation and with some broken turf (Thomas 1985b). The requirements and ecology of the butterfly in sand dune systems, which are used in some parts of the range are less well-studied. Here, they occupy mobile or grazed dunes and probably feed on Bird's-foot Trefoil, although the foodplant is not always clear (M. Oates pers. comm.). A summary of requirements is given in Appendix 2. 2.3 DISTRIBUTION AND POPULATION Distribution The Silver-studded Blue occurs across the temperate areas of Europe and Asia to Japan (Higgins & Riley 1970). In continental Europe, it occurs in a wider range of situations than in the British Isles, including alpine grassland, meadows, forest clearings and xerophytic shrubland. It occurs in most parts of Europe except northern Scandinavia (Higgins & Riley 1970). In Britain it was widely distributed at the turn of the century but it has since undergone a substantial decline and has become extinct except in southern and south-western England, and small parts of East Anglia, coastal Wales and the West Midlands (Heath et al. 1984, see Appendix 1). It is now common only in Dorset and Hampshire. The butterfly also occurs on the Scillies and Channel Islands. It used to occur in Scotland around the turn of the century (Heath et al 1984), although there is doubt over some of the records (Thomson 1980) (the only known specimen is from Loch Rannoch in 1936). It was once quite common in Cumbria and Lancashire (Birkett 1957) before its extinction in about 1942 (Thomas 1985b). Extinctions elsewhere in northern and central England and Wales occurred largely before 1940 (Thomas 1985b). 44% of pre-1940 10km squares were lost between 1945 and 1970 and 65% up to 1982 (Thomas 1985b, Warren et al. in prep). Population size The species tends to occur in high densities on small patches of suitable habitat. However, colonies can be short-lived, becoming extinct rapidly if habitat deteriorates. A survey in North Wales identified about 47 colonies, over 20% of which are large (>4000 adults at the peak of the flight period) (Thomas 1985b). Ten of these are on Great Orme's Head where 10 some 250,000 adults occur at the peak of the flight period; these populations appearing stable from 1983-1990 (Thomas 1993). Most patches occupied on limestone are between 0.1 and 8 ha and just one site is larger than 8 ha (Thomas 1985b). Heathland populations in North Wales (20) are smaller; the largest is at South Stack, Anglesey (c. 40,000 adults at peak); but most are 40-4000. 72% of heathland colonies occupy 0.1-1.45 ha and just one site is larger than 8 ha (Thomas 1985b). These also appeared stable in the period 1983-1990 (Thomas 1993). Sixteen colonies which have resulted from an introduction to the limestone of the Dulas valley are similar in size to the heathland population. The sole mossland colony has remained at 5000 adults on 4 ha (Thomas 1985b, 1993). In Suffolk, there are now just eleven colonies in three isolated regions. The species used to be abundant here and on the Brecklands further west (Mendel & Piotrowski 1985). Only one colony is substantial (c. 7000 adults at 1986 peak on 11 ha) and here numbers and the area occupied continues to decline (2000 on 6 ha in 1994) (Ravenscroft 1990a,b, 1994). All other colonies are small (<2000 adults on <2 ha) and less than 1% (19 ha) of available heathland was occupied in 1986 (13.5 ha in 1994). One small colony was founded between 1985-1994 and one became extinct. Two other colonies were established by introductions in 1986 (Ravenscroft 1992a,b). In Shropshire, one colony survives on 8 ha of Prees Heath (125 ha) (c 4000 adults at peak; Joy 1992). In Dorset, it is abundant on heathland (Thomas & Webb 1984), the subspecies cretaceus occupied 159 ha of limestone grassland in disused quarries on the Isle of Portland in 1985 (Warren 1986). The status of this unique subspecies is cause for concern: recent surveys suggest that many colonies are small and several have been lost in the last 15 years, primarily through increased quarrying (P. Green, pers. comm., A. Nicholson pers. comm.). In Devon, the butterfly occurs in 31 colonies on the East Devon Commons, but most colonies appear small and the majority of heathland is unoccupied (Thornett 1995). West Surrey supports 14 colonies of unknown size (Wilmott 1994). There is less detail available on the status of the Silver-studded Blue elsewhere in its range, such as in the New Forest, Cornwall, South Wales and Norfolk. Population structure The population structure of the Silver-studded Blue is one of the best studied of British butterflies. The species exists in metapopulations* and their function has been well defined in heathland populations in N. Wales (Thomas & Harrison 1992, Thomas 1996). Because suitable conditions on heathland are short-lived, it cannot persist on heathland that is: (1) not managed; or (2) too small to provide a continuation of conditions through natural events; or (3) too isolated to allow recolonisation should a population become extinct. The persistence of the Silver-studded Blue therefore depends on suitable habitat being continuously available within either relatively large areas of habitat or as a closely linked network of smaller sites. Within heathland metapopulations, local extinctions are usually * a metapopulation is a collection of local populations, connected by occasional dispersal, in which there are local extinctions and colonisations (Gilpin & Hanski 1991). 11 confined to small colonies of <0.9 ha (Thomas & Harrison 1992). On limestone habitats, extinctions and recolonisations most often occur on smaller patches (<0.2 ha) because conditions in these habitats usually persist for longer (Thomas & Harrison 1992). The recorded mobility of the Silver-studded Blue is low. In N. Wales, 89% of movements by adults within a day are <20 m, and none were recorded >50 m (Thomas 1985a). Isolated individuals sometimes move up to 400 m from a colony, but migration over 1km is thought to be very rare. Only about 1.4% of marked adults moved between habitat patches separated by 13-200 m in N. Wales (Lewis et al. 1995). No colonisation has been noted over 600 m in N. Wales, although all suitable patches of habitat within 1km existing populations tend to be occupied (Thomas & Harrison 1992). In Suffolk, 74% of adult movements were below 20 m, 7% >50 m and small pockets of adults occur within 500-600 m of colonies (Ravenscroft 1990a). However, larger scale movement does occasionally occur, an exchange of individuals having been noted between two colonies separated by 1.5 km and a colony was apparently founded over 3.5km in 1994 (Ravenscroft 1994). 2.4 LIMITING FACTORS Historical Habitat loss, through: (1) Destruction, mainly by agriculture, forestry, industry (including mineral extraction) and housing; (2) Decline of traditional management (turf and scrub cutting, burning and domestic grazing) leading to loss of early successional conditions and succession to scrub; (3) Decline in rabbit populations owing to myxomatosis, although recent rabbit recovery has reversed this in some areas. Ensuing effects on colonisation potential: (1) isolation of sites, preventing colonisation of suitable habitat; (2) reduction in rate of habitat creation owing to decline in traditional management; (3) reduction of site size, reducing likelihood of creation of suitable conditions sites through natural events in the absence of management. Current and potential Continuing destruction of heathland. Absence of suitable management. 12 within Inappropriate heathland management. Any decrease in rabbit populations through disease causing a reduction in grazing pressure (although many calcicolous grasslands are overgrazed). Associated effects of site isolation and heathland fragmentation as described above. Increased quarrying activities on the Isle of Portland. 2.5 RESUME OF CONSERVATION TO DATE The Silver-studded Blue has been well studied in many parts of its range, especially where the species is rare. It is regarded as a characteristic species of lowland heathland and an indicator of active lowland heathland management. Studies have been performed in North Wales (Thomas 1983, 1985a,b), East Anglia (Ravenscroft 1990a,b, 1994), Shropshire (Thomas 1985a,b, Joy 1992), Surrey (Wilmott 1994), Devon (Read 1985, Thornett 1995), Dorset (Webb & Thomas 1994), Sussex (Crane 1995) and parts of Dorset (Warren 1986), although not all of these describe status. There has been comparatively little study of status where the species is more common on the heaths of southern England e.g. in Hampshire and Cornwall. The Silver-studded Blue is well-represented on nature reserves, SSSIs and other protected areas owing to the high degree of protection afforded to heathland. In N. Wales, 17 of 47 colonies are on nature reserves (mainly LNRs on Great Orme's Head which is a Country Park, six on RSPB reserves, two on NNRs) and the remainder are privately owned (Thomas 1985b). In Suffolk, small populations occur on two NNRs, three on an RSPB reserve, one on NT land and most of the remaining sites are on SSSIs and Council owned (Ravenscroft 1994). In Surrey, parts of three colonies occur on NT ground and two on reserves of the Surrey Wildlife Trust. Only two sites are privately owned, the remainder are owned by various local Government authorities (Wilmott 1994). Elsewhere, large areas of heathland supporting the Silver-studded Blue are owned or managed by conservation organisations (e.g. in Dorset there are six NNRs and three RSPB reserves with the species and many smaller nature reserves of the Dorset Wildlife Trust and Local Authorities, A. Nicholson pers. comm.). On the Isle of Portland most colonies of the cretaceus race are within an SSSI and one colony occurs on a Butterfly Conservation reserve. The butterfly occurs on at least two Sussex Wildlife Trust reserves. In Hampshire, the Silver-studded Blue is ubiquitous in the New Forest, most of which is SSSI and managed by the Forestry Commission. Elsewhere in the county, Hampshire Wildlife Trust have at least three reserves supporting the butterfly. The RSPB manage Aylesbeare in Devon and Grange Heaths in Dorset, and in Cornwall there are large areas of NT ground e.g. at Holywell Dunes and on the Lizard, and at least two Cornwall Wildlife Trust Reserves with the species. In South Wales, the Silver-studded Blue occurs on at least one area of NT land in Pembrokeshire. In some parts of the range of the Silver-studded Blue there are specific heathland projects. These include:- the Heathlands Project Team of the RSPB in Dorset; English Nature's Wildlife Enhancement Scheme also in Dorset as well as Hampshire; the Suffolk Coast and 13 Heaths Project (Suffolk County Council); The Sandlings Project (Suffolk Wildlife Trust); the NE Hampshire Heathlands Project (mainly Local Authorities); the Surrey Heathland Project (mainly Local Authorities); and in Sussex, the Heathlands Project of the South Downs Conservation Board. Cornwall Wildlife Trust have a heathland conservation project in conjunction with the RSPB (NB. this is not an exhaustive list). The distribution and sizes of populations occurring in some parts of the range are well monitored (North Wales; Thomas 1983, 1985a,b, 1993, Thomas & Harrison 1992: Suffolk; Ravenscroft 1986, 1987, 1990b, 1994). Many other colonies are monitored by site managers, especially on nature reserves. These include wardens of the RSPB at Minsmere (Suffolk), Aylsebeare (Devon), Arne (Dorset) and South Stack (N.Wales) and English Nature at Studland, Hartland Moor and Stoborough Heath (Dorset). Branches of Butterfly Conservation (particularly Hampshire, Devon, Sussex, Surrey, Suffolk and Dorset) are active in monitoring populations and managing further sites. A full review of the status of the Silver-studded Blue on conservation sites (and their management priorities) and monitoring activities throughout England and Wales is required. Introductions A number of introductions and re-introductions of the Silver-studded Blue have been attempted by conservationists and justified because: (1) its habitat is now so fragmented; (2) the species has limited colonisation potential; (3) many suitable, unoccupied patches of habitat exist; (4) the suitability of these patches is relatively easy to assess. Re-introductions represent an attempt to restore dispersal in modern fragmented landscapes (Thomas 1993). Well-documented introductions of the Silver-studded Blue have been made in N. Wales and Suffolk and demonstrate the potential with this species. In the former, 90 adults were released in 1942 in the Dulas valley, near Great Orme's Head (Marchant 1956). There are now an estimated 16 colonies and 30,000 adults (at peak) spread along the limestone valley (Thomas 1985b, 1993). Six other introductions were made elsewhere in N. Wales 1978-1983: two are still extant (one heath, one calcicolous); two survived for 2-7 years (both heathland); and two failed (one calcicolous, one heathland) (Thomas & Harrison 1992). In Suffolk, introductions were made at two sites in 1986 using stock from a site due for development (Ravenscroft 1987). Both were extant in 1994 and one has become the largest colony in Suffolk and spread to nearby heathland (Ravenscroft 1992a,b, 1994). In 1994, stock from Prees Heath in Shropshire were released on the Wirral where the species used to occur (J. Joy pers. comm.). Management Management for the butterfly has been summarised recently by Joy (1995). Regional accounts and specific recommendations can be found in Thomas (1983, 1985b), Read (1985), 14 Ravenscroft (1990a,b, 1994), Crane (1995), Joy (1992) and Wilmott (1994). A summary of the main points is given in Appendix 2. Part 3 Actions and Work Programme This section has been divided into the standard headings Policy and Legislative; Site Safeguard and Acquisition; Land Management; Species Protection and Licensing; Advisory; International; Future Research and Monitoring; Communications and Publicity; Review. Actions appear under one heading according to their major role and/or aim, and are given a low, medium or high priority. The lead organisation(s) concerned for each action is/are named. Lead organisations concerned 3.1 POLICY AND LEGISLATIVE Action 1 PRIORITY: HIGH Include habitat requirements of the Silver-studded Blue where appropriate in management prescriptions for ESAs, Countryside Stewardship, Habitat Scheme and other schemes (Wildlife Enhancement Scheme etc.) MAFF, WOAD, EN, FA, CCW 3.2 SITE SAFEGUARD AND ACQUISITION Action 2 PRIORITY: HIGH Protect strategic metapopulations through management agreements and/or reserve acquisition. Action 3 PRIORITY: MEDIUM Ensure development proposals do not adversely affect existing or potential habitats. 3.3 ALL LAND MANAGEMENT 15 ALL Action 4 PRIORITY: VERY HIGH Promote beneficial heathland management in areas where the Silverstudded Blue exists, and restore links between populations. Action 5 PRIORITY: HIGH Promote beneficial grassland management for the calcicolous populations of Great Orme's Head and the Isle of Portland. Action 6 BC, WTs, FC(New Forest), FE, NT, EN, CCW, RSPB, LAs BC, LAs, EN, CCW. PRIORITY: MEDIUM Restore suitable habitats in former range, initially concentrating on areas close to existing populations (south-east, south-west, eastern and midland England, see Appendix 1). ALL 3.4 SPECIES MANAGEMENT, PROTECTION AND LICENSING Action 7 PRIORITY: LOW Conduct strategic re-introductions to suitable networks of habitats within former range using appropriate stock. Action 8 BC, EN, CCW, WTs, NT, RSPB. PRIORITY: HIGH Prepare separate action plans for high priority regions (North Wales, South Wales, Suffolk, Norfolk, Cornwall), following targets of UK Biodiversity Action Plan (within three years). BC, EN, CCW 3.5 ADVISORY Action 9 PRIORITY: HIGH Advise conservation agencies, site owners/managers and project officers of the relevant agri-environmental schemes and their role in promoting suitable habitat management for the Silver-studded Blue. 3.6 INTERNATIONAL No actions proposed 16 BC, EN, CCW, MAFF, WOAD 3.7 FUTURE, RESEARCH, SURVEY AND MONITORING Action 10 PRIORITY: HIGH Survey status of Silver-studded Blue in less well-documented parts of its range (South Wales, Norfolk, Cornwall and parts of Hampshire). Action 11 PRIORITY: HIGH Review the status of the Silver-studded Blue throughout its range, especially on sites of conservation activity and current or planned management. Action 12 BC, CCW, EN PRIORITY: HIGH Investigate ant habitat preferences on heathland and those of the Silverstudded Blue on southern heathland and sand dunes and the implications for habitat management. 3.8 EN, CCW, BC PRIORITY: MEDIUM Collate transect and management data from all monitored sites (in addition to BMS) annually and calculate annual index to compare trends on individual sites. Review and extend work if necessary. Action 14 BC, EN, CCW PRIORITY: MEDIUM Survey suitability of potential networks of sites throughout former range of the Silver-studded Blue and identify strategic sites for restoration and/or introductions. Action 13 BC, NT, WTs, CCW, EN BC, EN, CCW, ITE, Univs COMMUNICATIONS AND PUBLICITY Action 15 PRIORITY: HIGH Publicise this Action Plan, the status of the Silver-studded Blue and the measures being taken to conserve it. 17 BC, EN, CCW Action 16 PRIORITY: HIGH Produce management guidelines for use by staff preparing management plans for sites with colonies being entered into agri-environmental schemes. 3.9 BC, EN, CCW, MAFF, WOAD REVIEW Action 17 PRIORITY: MEDIUM Monitor this Action Plan annually and update every five years as necessary. BC, EN, CCW Key to abbreviations BC CCW EN ITE FA FC FE LAs MAFF NT RSPB Univs WOAD WTs Butterfly Conservation Countryside Council for Wales English Nature Institute of Terrestrial Ecology Forestry Authority Forestry Commission (New Forest) Forest Enterprise Local Authorities Ministry of Agriculture, Fisheries and Food National Trust Royal Society for the Protection of Birds Universities Welsh Office Agriculture Department The Wildlife Trusts 18 References Birkett, N.L. (1957). Insects. The Changing Scene, 1, 40-46. Penrith and District and Kendall Natural History Societies. Crane, R. (1995). Iping and Stedham Local Nature Reserve. Report on the Silver-studded Blue butterfly. Report to LNR Management Committee, Sussex. Dennis, R.L.H. (1977). The British Butterflies. Classey. Emmet, A.M. and Heath, J. (1990). The Butterflies of Great Britain and Ireland. Harley, Colchester. Fiedler, K. (1989). European and North-west African Lycaenidae (Lepidoptera) and their associations with ants. Journal of Research on the Lepidoptera, 28, 239-257. Fiedler, K. and Saam, C. (1995). Ants benefit from attending facultatively myrmecophilous Lycaenidae caterpillars: evidence from a survival study. Oecologia, 104, 316-322. Gilpin, M. and Hanski, I. (eds) (1991). Metapopulation dynamics: empirical and theoretical investigations. Academic Press, London. Heath, J., Pollard, E. and Thomas, J.A. (1984). Atlas of Butterflies in Britain and Ireland. Viking, Harmondsworth. Higgins, L.G. and Riley, N.D. (1970). A Field Guide to the Butterflies of Britain and Ireland. Collins, London. Jordano, D. and Thomas, C.D. (1992). Specificity of an ant-lycaenid interaction. Oecologia, 91, 431-438. Jordano, D., Rodriguez, J., Thomas, C.D. and Fernandez Haeger, J. (1992). The distribution and density of a lycaenid butterfly in relation to Lasius ants. Oecologia, 91, 439446. Joy, J. (1992). The conservation of the Silver-studded Blue butterflies (Plebejus argus) of Shropshire. Unpublished report to World Wide Fund for Nature (UK). Joy, J. (1995). Heathland management for the Silver-studded Blue butterfly. English Nature. Lewis, O. T., Thomas, C.D., Hill, J.K., Brookes, M. I., Crane, T.P.R., Graneau, Y., Mallet, J.L.B. and Rose, C.B. (1995). Dispersal and habitat tracking in metapopulations of a sedentary butterfly, Plebejus argus. Unpublished ms, University of Leeds. 19 Malicky, H. (1969). Verusch einer Analyse der ökologischen Beziehungen zwischen Lycaeniden (Lepidoptera) und Formiciden (Hymenoptera). Tijdschrift voor Entomologie, 112, 213-298. Marchant, A.J. (1956). Plebejus argus caernensis Thompson at Rhyd-y-Foel and Llanddulas. Entomologist, 89, 235. Mendel, H. and Piotrowski, S. (1985). The Butterflies of Suffolk. Suffolk Naturalists' Society, Ipswich. Mendel, H. and Parsons, E. (1987). Observations on the life history of the silver-studded blue, Plebejus argus L.. Transactions of the Suffolk Naturalists' Society, 23, 208. Nature Conservancy Council (1989). Guidelines for the selection of biological SSSIs. Nature Conservancy Council, Peterborough. Pierce, N. E., Nash, D. R., Baylis, M. and Carper, E. R. (1987). The costs and benefits of co-operation for the Australian lycaenid butterfly, Jalmenus evagoras, and its attendant ants. Behavioural Ecology and Sociobiology, 21, 237-248. Ravenscroft, N.O.M. (1986). An investigation into the distribution and ecology of the silver-studded blue butterfly (Plebejus argus L.) in Suffolk. Unpublished report to Suffolk Trust for Nature Conservation and Nature Conservancy Council. Ravenscroft, N.O.M. (1987). Changes in status of Plebejus argus colonies in Suffolk 19851987. Unpublished report to Suffolk Trust for Nature Conservation. Ravenscroft, N.O.M. (1990a). The ecology and conservation of the silver-studded blue butterfly (Plebejus argus L.) on the Sandlings of East Anglia, England. Biological Conservation, 53, 21-36. Ravenscroft, N.O.M. (1990b). The status of the silver-studded blue butterfly on the Sandlings, Suffolk 1990. Unpublished report to Suffolk Wildlife Trust. Ravenscroft, N.O.M. (1992a). The use of introductions for the conservation of a fragmented population of the silver-studded blue butterfly Plebejus argus L. in eastern England. In: Future of Butterflies in Europe: Strategies for Survival. Ed. by T.Pavlicek-van Beek, A.H.Ovaa and J.G. van der Made, pp. 213-221. Wageningen. Ravenscroft, N.O.M. (1992b). The fortunes of the silver-studded blue Plebejus argus (L.) (Lepidoptera: Lycaenidae) at artificial sites on the Sandlings, Suffolk. Entomologist's Gazette, 43, 157-161. Ravenscroft, N.O.M. (1994). Silver-studded Blue Butterfly: status, management and recovery 1994. Unpublished report to English Nature (East Region). Read, M. (1985). The Silver-studded Blue Conservation Report. Unpublished report to Joint Committee for the Conservation of British Insects. 20 Thomas, C.D. (1983). The ecology and status of Plebejus argus L. in North West Britain. Unpublished M.Sc. thesis, University of Wales, Bangor. Thomas, C.D. (1985a). Specialisations and polyphagy of Plebejus argus (Lepidoptera: Lycaenidae) in North Wales. Ecological Entomology, 10, 325-340. Thomas, C.D. (1985b). The status and conservation of the butterfly Plebejus argus L. (Lepidoptera: Lycaenidae) in North West Britain. Biological Conservation, 33, 29-51. Thomas, C.D. (1993). The Silver-studded Blue, Plebejus argus L.. In: Conservation Biology of Lycaenidae (Butterflies) (ed. T.R. New), pp. 97-99. IUCN Species Survival Commission. Thomas, C.D. (1996). Essential ingredients of real metapopulations, exemplified by the butterfly Plebejus argus. In: Aspects of the Genesis and Maintenance of Biological Diversity (eds. M.Hochberg, J.Clobert & R.Barbault), pp. 292-307. Oxford University Press. Thomas, C.D. and Harrison, S. (1992). Spatial dynamics of a patchily distributed butterfly species. Journal of Animal Ecology, 61, 437-446. Thomas, J.A. and Webb, N. R. (1984). The Butterflies of Dorset. Dorset Natural History and Archaeological Society, Dorchester. Thomson, G. (1980). The Butterflies of Scotland. Croom Helm, London. Thornett, R. (1995). Silver-studded Blue butterfly on the East Devon Commons 19881994. Report to Devon Branch of Butterfly Conservation. Warren, M. S. (1986). The status of the cretaceous race of the silver-studded blue butterfly, Plebejus argus L., on the Isle of Portland. Proceedings of the Dorset Natural History and Archaeological Society, 108, 153-155. Webb, N. R. and Thomas, J. A. (1994). Conserving insect habitats in heathland biotopes: a question of scale. In: Large-scale Ecology and Conservation Biology (eds P. J. Edwards, R. May and N. R. Webb), pp. 129-151. Blackwell, Oxford. Wilmott, K. J. (1994). The status and habitat requirements of the silver-studded blue butterfly in the county of Surrey. Unpublished report to English Nature. 21 Appendix 1 Distribution of the Silver-studded Blue in the UK (produced with kind permission of the Biological Records Centre). 22 Appendix 2 Heathland management for the Silver-studded Blue Heathland management for the butterfly has been summarised recently by Joy (1995). Regional accounts and specific recommendations can be found in Thomas (1983, 1985b), Read (1985), Ravenscroft (1990a,b, 1994), Crane (1995), Joy (1992) and Wilmott (1994). Grassland management for calcicolous populations is described in section 2.2 (see also Thomas 1985b and Warren 1986). Successful management aims to produce a mosaic of heathland of different stages and a continual presence of early successional vegetation. This is less important on large expanses of habitat where suitable conditions are likely to occur through random events such as disturbance and burns, and, if resources are limiting, management should focus on small heathland habitats, especially those linking larger populations. Suitable management techniques will vary from region to region and should be determined locally in association with management of the whole ecosystem. There are potential conflicts in management priorities on heathlands; suitable conditions for some uncommon birds and reptiles tend to be the later successional stages of heathland. Burning/grazing Periodic, patchy/small scale burning can be beneficial but needs to be considered carefully because of the risks involved. Summer burning is illegal (although can occur with a special MAFF/WOAD licence). However, accidental summer burns can be beneficial as they remove more of the litter layer than winter burns and promote recolonisation of heathers of open ground. All colonies in Suffolk show some evidence of previous burns and small summer burns are readily colonised (Ravenscroft 1994). Some colonies in Surrey are on accidental burns (Wilmott 1994). Large areas of sandy soils dominated by gorse would become suitable if burned; even isolated cutting and burning in small piles would produce patches of habitat. Conditions in most areas take 2-5 years to become suitable and remain suitable for 10-15 years. Moderate grazing by sheep and goats will maintain suitable conditions on calcicolous habitats indefinitely (Thomas 1985b) and rabbit grazing and digging will prolong conditions on heathland (Thomas 1985b, Ravenscroft 1990a, Joy 1995). The New Forest remains grazed. The presence of large herbivores may also help maintain soil disturbance. Re-instating grazing alone, however, is unlikely to be sufficient to restore conditions for the Silver-studded Blue. Soil disturbance and stripping Disturbance of the ground, especially if the topsoil is removed, is the main progenitor of suitable conditions in North Wales (Thomas 1985b). Most of the best sites here have had their topsoil removed at some time. Elsewhere, colonies are found in abandoned quarries (Dorset, calcicolous sites), old sandpits, disused sawmills (Suffolk), abandoned military installations (Shropshire, Suffolk) and firebreaks across heaths (most areas). Rotavation of sandy soils will re-create conditions, but may not suffice on more nutrient-rich soils (Thomas 1985b). Stripping topsoil may be a valuable tool in the restoration of former sites, after the 23 removal of any scrub and Bracken. Experimental strips removed on a colony being overwhelmed by grasses in Suffolk in 1986 are now the main parts of the site occupied (Ravenscroft 1994). Cutting/mowing Cutting and mowing can maintain areas for Silver-studded Blues in the absence of grazing, by keeping vegetation short and exposing areas of open ground. Small numbers of the butterfly frequent the edges of firebreaks on heathland that are regularly mown in many parts of the range (Thomas 1985b, Ravenscroft 1990a, Joy 1995), but this is not thought to represent good long-term management in Sussex (Crane 1995). However, in Dorset some large colonies occur in and around regularly mown heathland. Forage harvesting of mature heather areas can produce suitable habitat if debris is removed. 24