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J Insect Conserv DOI 10.

1007/s10841-007-9112-7

SHORT COMMUNICATION

The importance of resource databanks for conserving insects: a buttery biology perspective
Roger L. H. Dennis Peter B. Hardy Tim G. Shreeve

Received: 5 April 2007 / Accepted: 19 August 2007 Springer Science+Business Media B.V. 2007

Databases have become critically important in many areas of biological research. For example, searcheable nucleotide and protein sequence databases (e.g. Genbank, http://www. ncbi.nlm.nih.gov/Genbank/index.htm and Uniprot, http:// www.ebi.uniprot.org/index.shtml) and gene expression data (e.g. ArrayExpress, http://www.ebi.ac.uk/arrayexpress/ #ae-main[0]) hold important information to assist understanding of biological functioning at the cellular and subcellular level, and can be used to interpret evolutionary developments within a taxonomic group. Such databases are essential facilities for understanding complex processes and predicting interactions and systems dynamics. At the level of populations and communities, databases are available to predict agricultural pest infestation levels in relation to degree-days (e.g. UC IPM Online, http://www. ipm.ucdavis.edu/) and freshwater community composition in relation to physico-chemical condition (e.g. RIVPACS, http://www.ceh.ac.uk/sections/re/RIVPACS.html). Other databases (e.g. UK Buttery Monitoring Scheme, http:// www.ukbms.org/) have been used to identify phenological

R. L. H. Dennis (&) NERC Centre for Ecology and Hydrology, Monks Wood, Abbots Ripton, Huntingdon, Cambridgeshire PE28 2LS, UK e-mail: rlhdennis@aol.com R. L. H. Dennis Institute for Environment, Sustainability and Regeneration, Staffordshire University, Mellor Building, College Road, Stoke on Trent ST4 2DE, UK P. B. Hardy 1 Winstanley Road, Sale, Cheshire M33 2AT, UK T. G. Shreeve School of Life Sciences, Oxford Brookes University, Headington, Oxford OX3 0BP, UK

changes (e.g. Roy and Sparks 2000) and changes of abundance (Roy et al. 2001; Thomas et al. 2004; Thomas 2005). With few exceptions (e.g. Syrph-the-Net, http:// www.iol.ie/*millweb/syrph/syrphid.htm), those databases available for insect conservation mostly comprise species lists, locations and imprecise descriptions of vegetation communites; their use in predicting spatial and temporal occurrences is more limited but in specic circumstances they can be used within metapopulation modelling processes (e.g. Gutierrez et al. 1999; Davies et al. 2005, 2006). Such data are now being used via the focal species approach within exercises in landscape design and restoration, though the approach is not guaranteed of success (see Lindenmeyer et al. 2002). If molecular data can be used predictively then there is every chance that correctly designed and inclusive databases can be used to aid insect conservation. Both subcellular and population/community processes are highly complex and there are parallels in hierarchical organisation of data between molecular based systems and whole organism based systems (Fig. 1). It is logical to argue that a correctly designed database can be as useful for conservation as molecular databases are for understanding molecular and cellular processes. Here, we develop our arguments about the applicability of appropriate databases using butteries as a model taxon. This group is perhaps the best known group of insects in the world; they are heralded as regional indicator taxa for environmental changes (Thomas 2005; Thomas et al. 2004).

Why is a database needed? For a group of insects allegedly the best studied in the world, it may come as a surprise to those not working on

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J Insect Conserv Fig. 1 Parallels between (A) molecular, cellular and organism traits/processes and interactions supported by fundamental genomic information and (B) the spatial and temporal persistence of populations, based on major trait/processes and their interactions supported by fundamental information about resources, life history and behaviour. The major box in A represents within-individual processes and in B processes that contribute to the spatial occurrence and persistence of individuals and populations

A
Gene sequence

Selection Metabolic control systems Regulation

Functional individual

Cell signalling Protein sequence Protein structure

Growth and development

Available consumable resources

Utilities and substrates

Gene sequence

Selection Population and community level interactions Functional individual Regulation Control of resource use Growth, development and morphology

Potential life history, resource use and behaviour

Local life history and local resource use

Realised behaviour Available consumable resources

Utilities and substrates

them, and when so much is claimed for them, that so little is known. Much useful data can be gleaned on individual species from numerous texts (e.g. Emmet and Heath 1991; Dennis 1992, 1993; Asher et al. 2001), regional atlases and from autecological studies, many of which remain unpublished. But, these data are incomplete, often imprecise on key resource requirements, and lacking for more ubiquitous species which escape conservation concern. Effective conservation is hampered by this lack of information. Whilst the requirements of a few species, which have been the focus of intensive research, are relatively well known we have little information on the remainder. Thus the effect of specic conservation measures for a few species on the remainder cannot be predicted. When land areas are relatively undeveloped and large enough, landscape and resource heterogeneity are sufcient to maintain species and biodiversity, since different species require different resources. At a regional scale ([104 km2), resource turnover via geomorphic processes and vegetation dynamics is sufcient to maintain heterogeneity and therefore species persistence. However, in smaller areas and large regions long exposed to human development, the persistence of resources becomes increasingly uncertain and as natural and seminatural areas become smaller and scarcer there is an increasing need to have information on organisms habitats and their distributions, to know what is needed and what may be missing. We argue that there is sound evidence to indicate that current approaches to buttery conservation are not wholly effective and alternative

approaches are necessary to arrest and reverse biodiversity decline. Butteries clearly fall into the category of a agship taxon but they are not indicators of all environmental and insect changes (JICO Special Issue 2 on Habitats 2006). They are indicators of disturbing losses of resources within seminatural biotopes, difcult to ascertain from surveys of biotope cover (Thomas et al. 2004; Thomas 2005), and of climatic change (Dennis 1993; Thomas et al. 2001; Hill et al. 2002; Franco et al. 2006). For this group, distribution losses over the past decade or two appear to have been little short of spectacular (Van Swaay and Warren 1999; Fox et al. 2006, 2007). With few exceptions this trend continues despite management based on metapopulation models for spatial dynamics of selective species (Hanski and Gilpin 1997; Ehrlich and Hanski 2004) or on the basis of the maintenance of biotope or specic vegetation types. More recently, these approaches have been challenged as being too simplistic (Dennis et al. 2003, 2006, 2007), with the suggestion that vegetation type and/or biotope based denitions of habitat are inappropriate for buttery conservation. Habitat patch numbers, areas and isolation are obviously basic to an organisms spatial integrity and thus its conservation, but insufcient in themselves for achieving long-term persistence, particularly if based on imprecise (vegetation class type) denitions of habitat patches. European legislation has also focused conservation effort on priority species and habitats (=biotopes and vegetation units). However, whilst this focus may facilitate

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the conservation of particular species and some specic vegetation associations, the general trend for butteries is still one of overall decline. In some EU member states the rates of decline of non-listed species are far greater than those of listed species (Van Swaay and Warren 1999; Fox et al. 2006, 2007). Priority species are those which use restricted resources in current landscapes and the key question has to be whether a focus on such species will facilitate the conservation and enhancement of other species. A tacit assumption is that this will be the case. However this may not be so. Ideally, focal, or umbrella species should be selected on precise criteria (Sanderson et al. 2002; Coppolillo et al. 2004) and to be truly representative of a range of species, the traits and resources of the selected species need to be compared with those of other species. Thus, the correct selection of focal species dictates that all the attributes and resource requirements of associated species are known. Therefore, there is a requirement for detailed knowledge of species tolerances and resource requirements, in effect a need for a database containing relevant information. Only once such a database has been developed can the representativeness of focal or priority species be gauged. However, once the data are available, the need for focal species for conservation science has gone, although they may act as a vehicle for campaigning or publicity. A comprehensive database can also serve as a powerful tool in predictive ecological studies at the multi-species level and facilitate action to redress the decline of species. The use of appropriate data in predicting species distributions will also be of growing importance with increasing anthropogenic forcing of climate change and environmental degradation. There is a strong analogy between conservation biology and molecular based approaches to understanding organism function and persistence (see Fig. 1). The latter requires the understanding and inclusion of a number of traits and processes in predictive modelling for any part of the pathway. At the core of all these processes is functional genomics. The ultimate goal of conservation biology has to be to maintain and enhance populations of species and biodiversity. In effect, if the reasons why a particular organism occurs in a particular location are not understood, and only a limited part of the pathway is examined, there is high probability of failure. At the core of this has to be an understanding of resource requirements and the complexity of trait/process interactions. We argue that there is a pressing need for fundamental data on resource requirements; it is the equivalent of genomic data in molecular based studies. What should constitute the database? Fundamental to conservation is an understanding of what determines where species exist, what determines their

persistence and their mobility. For butteries there is a focus on the habitat as the determinant of where species occur, but there are important issues about how this should be dened. Recently, advances have been made in understanding how habitats should be dened (Dennis et al. 2003, 2006). An organisms habitat is most logically dened by the spatial occurrence of resources in the right condition. This resource-based denition of habitat (Dennis et al. 2003; Shreeve et al. 2004) contrasts with the traditional view of interpreting habitat as vegetation units or biotopes (Dennis et al. 2006, 2007) which demonstrably fails to encompass true habitats of organisms such as insects (Dennis 2004a, b). For butteries, resources incorporate more than host plants in the right condition. It encompasses all the resources required for all activities of the adult stage (e.g. nutrient sources, basking sites, roosting sites) and for all the developmental stages. The availability of all required resources, in the right context and condition for individuals to locate and use them, therefore denes the habitat. Understanding resource requirements, should, in theory, assist conserving organisms since it more closely denes the habitat than does a vegetation association based denition of habitat. We argue that a resource-based database is required to facilitate effective conservation, in particular: to understand the potential for species in different land-use units, to understand the reason why species that can co-occur respond differently to environmental change and land unit management, and to predict how species may repond to future environmental change. A resource-based approach to the database will enable species exact requirements to be described and be preferable to a vegetation association based data set which is an imprecise surrogate method for determining species requirements. Further evidence for the need to move to a resource-based approach and away from a vegetation association/biotope approach is provided by species co-existing in one location (biotope) but not in another and showing different responses to management regimes (Dennis et al. 2003; Shreeve et al. 2004). Several fundamentals of the resource-based approach to dening the habitat are now supported. First, buttery species require a range of resources, not just larval host plants, but other consumer resources (i.e. nectar, sap; Tudor et al. 2004; Hardy et al. 2007) and an array of utility resources. These include structures for a range of activites including roosting, thermoregulating and mate location (e.g. Plebejus argus, Dennis 2004a; Dennis and Sparks 2006; Hipparchia semele, Maes et al. 2006; Callophrys rubi, Vanreusel and Van Dyck 2007). Regarding adult consumer resources a number of butteries depend heavily on substrates other than nectar owers as adults (e.g. Apatura iris 95%; Neozephyrus quercus 53%; Limenitis camilla 31%) (Hardy et al. 2007). Second, quality of

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resources is a vital issue. It has long been known that rare buttery species use but a fraction of an apparent available resource, including larval host plants (e.g. Maculinea arion, Thomas 1984). This is also the case with common butteries; for example, nettle patches of suitable quality and size for Inachis io and Aglais urticae can occupy as little as 8% of the total number of patches (Dennis 2007). It is inevitable that resource quality is also supported in a metapopulation context (Dennis and Eales 1997, 1999; Thomas et al. 2001). Third, conditions matter, particularly local climate; unquestionably they form critical resources. Butteries are ectotherms and highly dependent on ambient conditions and sunlight; local climate and microclimate is a resource bound up with topography and vegetation structure (Shreeve 1992; Dennis 1993; Bryant and Shreeve 2002). A classic case is the restriction of early stages of Limenitis camilla to Lonicera in partial shade because the light environment affects the ability of larvae to construct their intricate feeding platforms (Fox 2005). Fourth, resources abound in what has been labelled the matrix; species regarded as dependent on habitat patchworks nd and use resources within the matrix (e.g. Maniola jurtina and Pyronia tithonus; Dennis 2004b) and species search the matrix for resources, often nding them, and regularly using resources in this context (e.g. Pieris species; Dennis and Hardy 2007). Some resources are profoundly embedded in the matrix in unexpected contexts; for instance, hill tops used by Lasiommata megera for mate location at elevations unlikely to support overwintering populations (Dennis and Dennis 2006, 2007). Having sound resource data lies at the foundation of another division in conservation, that of species into generalists and specialists (Dennis et al. 2004). Where the division is carried out on limited criteria (i.e. host plant number; host plant abundance, biotope type, biotope structure; Asher et al. 2001; Fox et al. 2006) without reference to the array of resources making up habitats used by buttery species, there is a risk of producing erroneous binary classications of species, and aberrant indications of trends. Where more criteria are used, species can be divided further into functional groups (Shreeve et al. 2001), reecting generalists, ruderal species, more specialised species associated with open xeric environments or species associated with canopy forest structures or features that currently occur in woodland, but not specically dependent on tree structures. Since species of different functional groups may co-occur within particular biotopes when their resources coincide it clearly indicates that a resource-based approach is essential. Species of different functional groups react differently to changes of resource circumstances, but on the basis of biotope asociation there would be no indication of differential responses. Furthermore, reliance on biotope and a limited set of resources may lead to

underestimations of where species can occur. For example, Argynnis adippe was once described as widespread and common within much of Britain (South 1906; Frohawk 1934). In the southern part of its geographic range it still is ubiquitous. In the past, and currently in southern and central Europe it occurs in a range of woodland structures, but in the British Isles it is now restricted to a few open Pteridium dominated areas and limestone rock outcrops where woodland has been cleared (Buttery Conservation undated) for which specic management practices are advocated. The resources this buttery use are not just restricted to these land-use types and by focusing on current (limited) vegetation associations the potential of the landscape for this species is seriously underestimated. In addition a focus on just host plants and biotope has led to management regimes which may actually be detrimental. The buttery is reliant on open scrub and wood edge shrubs, these are key non-consumable resources for adult thermoregulation, mate location and day-time roosting. Management for this buttery actually discourages these essential resources.

What is the current state of the buttery resource data? It is clear that a proper basis for conserving butteries must come from a true understanding of their requirements. Currently, two datasets are being developed by us; one on consumer resources (larval host plants and adult feeding sources; www.geocities.com/pgll@btopenworld.com/ resources/resources.htm; PBH/RLHD) and the second on utilities and conditions (Oxford Brookes University; TGS) incorporating European as well as British species. The database on consumer resources has arisen out of inadequacies disclosed during research on buttery ecology (Dennis et al. 2000; 2004). Adult food resources, initially based on Porter et al. (1992) and Corbet (2000), were found to be sparse and incomplete. Data on nectar ower and alternative substrate use now extend to 11,098 records and 49,202 observations on 74 buttery species and 519 plant taxa as of April 1 2007; 11,016 records for the 60 butteries that breed in Britain. The host plant data comprise 5,595 records (5,321 for the breeding species). The adult feeding data have been accumulated from 267 sources, largely unpublished, whereas the host plant data have been extracted from 196 sources that are mainly published. The elds in the database for adult feeding are: date of record, (post-1974) county (Heath 1976), grid reference to 10 km2; buttery species and Bradley and Fletcher (1986) code (see Emmet and Heath 1991), plant species or substrate, plant BRC code and family (Stace 1997 number), number of individual butteries observed for each entry in the le or record (counts of individuals), and source of

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record. For larval host plants the elds are: Bradley and Fletcher (1986) code numbers (Emmet and Heath 1991), buttery species, plant species, plant family, reference source, and sub-reference to earlier source, type of record and summary status of plant as a host plant. Regarding type of record, each host plant record is coded up for a number of categories (E, egg laying, or eggs found in the eld; L, larva(e) found in the eld; N, successful breeding in natural conditions from egg to pupa; B, successful breeding in laboratory situation; F, failure to oviposit or develop successfully in laboratory situations; R, reference extracted from previous source). Host plant status, based on these records, is determined from the coding of records and designations in the literature into: M, main; S, subsidiary (auxiliary/secondary; generally regarded as insufcient without the presence of main host plants); N, novel (alien plant to the British Isles); U, unsuitable/unlikely host plant. X, denotes a likely mistake in oviposition or deliberate oviposition off the host plant (Wiklund 1984). The host plant is also categorised as to any regional limitation: GL, geographical limitation (locality/area/region specied). The second database developed out of study of natural ecological groupings amongst British butteries (Shreeve et al. 2001). Initially it included 136 binary state attributes coded up for 57 species within the British Isles (see Shreeve et al. 2001, Appendix 1 for attributes); these included an array of life history and behavioural traits through all life history stages. This is being built on and now includes 161 attributes, including microhabitat location, and is being extended to species of the European mainland. Incorporated into this le are a large number of ecological measures taken from previous works. This le is available from one of us (TGS). These les represent a substantial advance over data held but 5 years ago. Records now number more by orders of magnitude and for consumer records are increasing exponentially; a clear idea is held of the nectar owering plants used by individual species, the importance of other substrates used by adults for some species (e.g. Apatura iris; Limenits camilla; Hardy et al. 2007) and the status of host plants listed for species. Perhaps most importantly, a protocol has been established for collecting data on consumer resources, at last matching that of museum specimens of the insects themselves and with extension of the service to utilities the prospects are good for having a sound database on British butteries. However, at the moment, there are substantial deciencies in the data. First, for some species we have limited records for even key resources: 29 buttery species have less than 50 records for adult feeding (Hardy et al. 2007). For some utilities (e.g. resting sites and moulting sites of larvae), we have very limited information. Second, supporting data for records, including the origins of the records

and geographical locations, are missing, indicative of gaps in some of the comprehensive sources. Of the 11,098 adult feeding records, 4,345 lack counts of the number of individuals concerned. Generally, the host plants of British butteries are largely taken for granted. However, of the 5,595 records, 5,079 (4,843 for the 60 species breeding in Britain) convey no specic details for the type of record, whether a main, subsidiary or novel host plant, whether a record of egg laying, larval feeding or successful breeding in the wild or laboratory. This is a serious problem for larval grass feeders (Parker 2003; Dennis et al. 2005), but even for common butteries there are missing data. As an example, we have little idea just what plants are used by larvae of Pieris brassicae, and how successfully outside cabbage patches and away from calcareous cliffs (Dennis and Hardy 2006). New host plants are still being recorded for species (e.g. Phalaris arundinacea for Ochlodes venata; Smart 2006) and information for grass and sedge feeding species is undoubtedly lacking (Parker 2003). Third, there is currently spatial bias in the data. Some 4,180 adult feeding records (39.4%) derive from a single county (Greater Manchester); the next largest contributions (391 or 3.69% from Hereford and Worcester, 345 or 3.25% from Gloucestershire) are much smaller, some 3,621 records lack any county provenance, and there are only 19 records (0.18%) from Eire and 8 (0.07%) from Northern Ireland. Some 5,350 records lack grid references. Fourth, data on resource use lack qualication; the quality dimension is missing in observations. Information on microhabitat use, at the level of small patch descriptions, is lacking for the majority of species, but this should provide important information about the quality and context of the resources that are used. Butteries not only do not ll the distributions of their basic host plant resources (Dennis and Shreeve 1991), but only use a fraction of any basic resource such as a larval host plant at any site. This was recognised decades ago for Maculinea arionwhere decline to extinction continued in the face of abundant thymeand for Satyrium pruni and Thecla betulae, which only use a fraction of available Prunus spinosa (Thomas 1974, 1980). Finally, most of the data are not the products of site systematic surveys, and/or based on appropriate sampling protocols. Generally, there is no explicit reference to location, and thus to spatial variability, and we have little idea other than for species subject to more thorough autecological studies how the various resource attributes covary within sites. As conservation operates through management of sites, this is a major failing of data on buttery resources. There is one other area where there is currently a gap in information. Butteries are thermophilous and all stages are dependent on specic microclimates. Data for only a few species exist on basking site temperatures and solar

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radiation loads required for adult activity (e.g. Pararge aegeria, Shreeve 1984, 1985; Maniola jurtina, Maier 1998; Pieris napi, Wilcockson 2002; Polyommatus icarus, Howe 2004) and for egg-laying sites (e.g. Polyommatus bellargus, Roy and Thomas 2003, Hamearis lucina, Sparks et al. 1994); scant information is available for larval development in relation to thermal regimes (e.g. Euphydryas aurinia, Porter 1984; Inachis io, Aglais urticae, Polygonia c-album and Vanessa atalanta, Bryant et al. 2000). Yet, the thermal environment is a crucial determinant of survival and reproductive success, and has a critical role in determining whether a particular location is occupied or not. The analytical tools to relate thermal regimes to vegetation structures are in existence (e.g. Bryant and Shreeve 2002), but the required thermal regimes are largely unknown.

Future demands of a database. What are the priorities? For British butteries, there are two kinds of data. The rst is that discussed extensively above, consisting of a wide array of records being made independently by large numbers of observers on all aspects of buttery life history and behaviour in relation to substrates; much data of this kind are already published, but not fully accessed, in literature dating back over the past 150 years. Second, there is site based, species-specic data emanating from autecological studies; much of this exists in the form of research theses and reports on rarer species. Respectively, these two types of data can be conveniently referred to as the Biological Resources Database (BRD) and the Site Database (SD). These two types of data (databases) require different handling. First, the BRD: It has become evident with the accumulation of adult feeding and larval host plant data that many of the same recording elds apply. It is also becoming obvious that there is no reason why these same elds should not be applied to utility data. As the list of utility variables is now largely complete, there is no reason why quantitative data on these variables, presently in binary format for attributes on species, should not be collected in much the same way as nectar feeding records and host plant records. What is required to start the process in motion is a clear guide to the collection of resource data, to variability (attribute states) in utility variables and recognised coding for the attributes and substrates. The data would best be stored in a single web-based database to which the public have direct access and to which they can contribute their observations through a quality control lter. The data are easily acquired during standard atlas (BNM; Asher et al. 2001; Fox et al. 2006) and transect (BMS; Pollard and Yates 1993) recording. The technique of supplementing standard and synoptic surveys with behavioural and resource data has now been successfully

applied in trials for both transect recording (Dennis 2004b) and for atlas recording (Dennis and Hardy 2007; Hardy and Dennis 2007). The basic purpose is to collect data on what individual butteries are actually doing, on what and where, to provide a key to what they use and need and the problems they face; thus, individual records for butteries are minimally coded up for species, sex, behaviour, resource substrate and any other ancillary data (i.e. weather conditions, aspect, topography, management, biotope etc.) that assist interpretation of behaviour. It is particularly disappointing to think how much valuable information is not being recorded during atlas surveys or supplementary to transect recording. Currently, there is little information even on biotope use among British butteries. Atlas records for butteries relate butteries to squares and to the biotopes recorded in those squares, of which there may be a number, but not to specic biotopes coinciding for grid squares. British Buttery Monitoring Scheme transect sections are only currently in the process of being coded up for vegetation structure (Roy personal communication). Support for a BRD is substantial; the data provide the basis for more advanced dedicated survey and experiment, the latter continuing to supply invaluable information for conserving butteries in Britain (e.g. Polyommatus bellargus, Roy and Thomas 2003; Leptidea sinapis, Jeffcoate 2006). The SD is a very different construct; it provides the all-important conduit for spatially explicit data on a range of resource variables and attributes. To understand properly how species persist in different landscapes it is essential to know what resources they access and use successfully at a site scale. A current detractor is that objectives of site studies vary in their inclusiveness of the relevant biological data (behaviour in relation to resource use) on species studied. Currently, there is no inventory of site based studies which may comprise BSc and MSc dissertations as well as doctoral theses and reports from funded work by governmental and national, regional or local organisations (e.g. National Trust; Buttery Conservation; County Trusts) as well as private unpublished, unfunded studies. An inventory is clearly a rst step, followed by a metaanalysis of its contents, methods for which are now being advanced (Pullin and Knight 2001). The development of both databases should provide clear direction as to what data are missing on species and which species therefore require immediate attention in terms of data collection. For conserving butteries and other arthropods there is an urgent need, therefore, to acquire data on resource requirements. For many species, this is blatantly lacking. Yet, much of these data can be acquired easily enough through survey and experiment by students engaged in GCSE and A level projects in schools and BSc and MSc dissertations in colleges and universities as well as those currently engaged in a range of buttery surveys. The

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urgent request is that teachers and tutors encourage eldwork to obtain resource data on arthropods whatever their taxonomic inclination and make this available in the literature. Numerous journals exist for this very purpose; their editors keen to encourage contributions (e.g. UK Bulletin of the Amateur Entomologists Society; Entomologists Monthly Magazine) and entomologists of the future. In the face of climate change and land-use change, knowledge of species resource requirements is critical. Some species conned to specic, isolated, sites are thought potentially to face future extinction, yet we do not know their resource requirements and therefore the potential of the landscape to support them is largely unrecognised. A focus on resources and a resource-based habitat denition (Dennis et al. 2003, 2006, 2007; Shreeve et al. 2004), together with information on the distribution of resources within the landscape, may well clarify and resolve regional differences of spatial abundance and vegetation associations, and foster effective conservation effort. Data collection has its costs; particularly the easily forgotten time and effort freely given by volunteers, and tre as well as thus a database must have a sound raison de transparent procedures. In converting reasons for obtaining resource data into objectives, several are prominent. The rst, to conserve species in strict species-specic programmes, a complete understanding of resource requirements is critical for appropriate management; missing out essential resources courts failure in site management and only having partial data may result in the potential for future changes to be underestimated. The second follows on from this, comprising site auditing for numbers of species to assess whether sufcient and appropriate resources are available within sites. Inevitably, there will be shortcomings, such as insufcient resource space and deciencies in neighbourhood resource associations. Third, and perhaps most important in the face of climate change and landscape change, is that the resource data will provide the means for mending the matrix (Dennis and Hardy 2007), moving away from a binary view of landscape (patch versus matrix) and towards developing networks rather than patchworks of different resource types. This is where site-specic data are crucial because they provide the only means for understanding species population dynamics. By-products of this approach provide a counter to the wanton destruction of weeds and the ecologically motiveless manicuring of hedges and verges. Resource data collection provides essential corroboration of policies generated through use of indicator taxa where contrasts in patch and matrix become increasingly blurred with the widening focus on more species. If a cohort of species is being used as indicators for other organisms then it becomes essential to know what resources the indicator species actually represent. Finally,

we need better data to understand resource use by species. Admittedly, there is a pure scientic set of objectives to understand differences in species ecology, but the by-product of pure research is that it generates more data, more ideas and new methods. Databases are selfperpetuating as further research using them encourages enhancement, all of which should lead to more integrated conservation programmes and better management. The existing two databases have already been used to predict species co-occurrences and group species by strategies (Shreeve et al. 2001), and have contributed to understanding the links between buttery strategies and host plant strategies (Dennis et al. 2004) and between consumer strategies of adults and larvae (Dennis et al. 2005; Hardy et al. 2007). We advocate that our initial approaches to establishing a resource database for butteries primarily within the UK is extended to at least Europe and that it is used predictively or its powers of prediction be tested. We need more comprehensive data and urge that a mechanism is established to facilitate data acquisition. This could mirror mechanisms Genbank and its equivalents use to acquire sequence data when relevant material is published. In effect, a web based interactive database for conservation should be the equivalent of a genomic database for molecular based bioinformatics. References
Asher J, Warren M, Fox R, Harding P, Jeffcoate G, Jeffcoate S (2001) The Millennium Atlas of Butteries in Britain and Ireland. Oxford University Press, Oxford Bradley JD, Fletcher DS (1986) An indexed list of British butteries and moths. Orpington, Kent, 119 pp Bryant SR, Shreeve TG (2002) The use of articial neural networks in ecological analysis: estimating microhabitat temperature. Ecol Entomol 27:424432 Bryant SR, Thomas CD, Bale JS (2000) Thermal ecology of gregarious and solitary nettle-feeding nymphalid buttery larvae. Oecologia 122:110 Buttery Conservation (undated) http://www.buttery-conservation. org/conservation/defrafactsheets/butteries/high_brown_fritillary. pdf Coppolillo P, Gomez H, Maisels F, Wallace R (2004) Selection criteria for suites of landscape species as a basis for site-based conservation. Biol Conserv 115:419430 Corbet SA (2000) Buttery nectaring owers: buttery morphology and ower form. Entomol Exp Appl 96:289298 Davies ZG, Wilson RJ, Brereton TM, Thomas CD (2005) The reexpansion and improving status of the silver-spotted skipper buttery (Hesperia comma) in Britain: a metapopulation success story. Biol Conserv 124:189198 Davies ZG, Wilson RJ, Coles S, Thomas CD (2006) Changing habitat associations of a thermally constrained species, the silver-spotted skipper buttery, in response to climate warming. J Anim Ecol 75:247256 Dennis RLH (ed) (1992) The ecology of butteries in Britain. Oxford. Oxford University Press, Oxford

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