USE OF THE SYRPH THE NET DATABASE 2000

M.C.D.Speight, E.Castella & P. Obrdlik

SYRPH THE NET: THE DATABASE OF EUROPEAN SYRPHIDAE (DIPTERA)

Volume 25

Series Editors: Martin C.D.Speight, Emmanuel Castella, Petr Obrdlik & Stuart Ball USE OF THE SYRPH THE NET DATABASE 2000

M.C.D.Speight Research Branch, National Parks and Wildlife, 7 Ely Place, Dublin 2, Ireland

E.Castella, Laboratoire d'Ecologie et de Biologie Aquatique, Université de Genève, 18 chemin des Clochettes, CH - 1206 GENEVE, SWITZERLAND

P.Obrdlik WWF Auen Institut, Josefstraße 1, D-7550 RASTATT, GERMANY

Syrph the Net: the database of European Syrphidae (Diptera) Volume 25 Speight, M.C.D., Castella, E., Obrdlik, P. and Ball, S. (eds.) 2000 compilation of the Syrph the Net database received funding from: contract STEP/CT90/0084 (Science and Technology for Environmental Protection), European Commission

this publication may be referred to as:

Speight, M.C.D., Castella, E. & Obrdlik, P. (2000) Use of the Syrph the Net database 2000. In: Speight, M.C.D., Castella, E., Obrdlik, P. and Ball, S. (eds.) Syrph the Net, the database of European Syrphidae , vol.25, 99 pp., Syrph the Net publications, Dublin

ISSN 1393-4546 (Series)

Syrph the Net Publications Dublin 2000 © M.C.D.Speight 2000 USE OF THE SYRPH THE NET DATABASE 2000

CONTENTS

Preface

Chapter 1: INTRODUCTION

1.1. Invertebrates in environmental interpretation and evaluation 1.2. The basic approach adopted in the syrphid database 1.3. The ingredients of the predictive process 1.3.1. Regional lists 1.3.2. Syrphid Habitats 1.4. Modelling 1.5. Origin and development of the syrphid database

Chapter 2: FIELD PROCEDURES

2.1. Site description 2.1.1. Use of the Habitat Survey Form 2.2. Field-sampling procedures 2.2.1. Placing of Malaise traps 2.2.2. Sample collection and trap maintenance 2.2.3. Field sampling strategy: obtaining representative samples of site faunas 2.2.3.1. Choice of sampling period 2.2.3.2. Duration of field campaigns 2.2.4. Inventory survey

Chapter 3: LABORATORY PROCEDURES

3.1. Treatment of samples 3.2. Determination of sorted specimens 3.3. Recording the determined specimens

Chapter 4: DATA PROCESSING PROCEDURES

4.1. Field data files 4.2. The basic site interpretation procedure 4.3. Statistical/analytical techniques 4.3.1. Use of multivariate ordination techniques with the database 4.3.1.1. Reciprocal ordination of the species and sampling stations 4.3.1.2. Reciprocal ordination of the species and their attributes 4.3.1.3. Simultaneous ordination of two matrices

Chapter 5: APPLICATIONS OF THE DATABASE

5.1 Application of the FAEWE procedure for assessment of the “ecosystem maintenance” function of a site 5.1.1. The functional assessment procedure (FAP) 5.1.2. Salient features of the functional assessment procedure

5.2. Use of the database in general site management 5.2.1. Use of the database in site restoration 5.2.1.1. Replacement of missing ecosystem components 5.2.1.2. Replacement of one ecosystem by another

5.3. Comparisons between regional lists

Chapter 6. PROGRESS & LIMITATIONS

REFERENCES

Appendix 1. Habitat Survey Form

Appendix 2. Nomenclaturally complete species list

Appendix 3. Taxonomic literature: genera keyed out by major works PREFACE

By far the greater part of this volume has been culled from material already published by the authors (see Castella and Speight 1996, Castella et al 1994, Murphy et al 1994, Speight 1996b, Speight 1997, Speight and Castella 1995), though the published material has had to be augmented to provide a coherent picture. It is not intended as a manual for the interrogation of the syrphid database, but more as a collection of examples of potential usage, together with suggestions on how field, laboratory and interpretation work might be standardised to serve particular objectives. It is assumed that the potential user has a familiarity with the manipulation of Excel spreadsheets, or access to instruction manuals on their use. The first version of the database was produced for use in an EU funded research project which formed part of the STEP programme. This project, the “Functional Analysis of European Wetland Ecosystems” project, or FAEWE project, is referred to at various points in the present text.

1 USE OF THE SYRPH THE NET DATABASE 2000

Chapter 1. INTRODUCTION

The syrphid database has been set up for use as a tool in: a) environmental interpretation, b) site evaluation/management, c) the study of Syrphidae.

It comprises a series of spreadsheets and text files grouped into volumes, each of which deals with a particular topic. The topics covered so far are: Species accounts Macrohabitats Microsite features Traits Range and Status Use of the database

The Species Accounts volume is text throughout, but the other volumes each include both text and spreadsheet files, as is the case in the present volume, which is focused broadly on use of the database. Details of the coverage of each volume are given in an introductory text at the beginning of each volume, and are not repeated here. A readme file is provided to help the user associate the constituent files of each volume of the database correctly. More than 550 European syrphid species are now covered by the database, out of a total European fauna of c750 species.

The spreadsheets have been created, saved and used in an Excel™ spreadsheet environment. Excel has been used because it allows maximal flexibility of use of the files subsequently. Any user who has need to repeatedly interrogate the database in a specific way can convert the files into, for example, Microsoft Access files and construct whatever database management system best suits his/her requirements for speeding up the interrogation process.

The spreadsheets provide a digitised transcription of information available about the species considered. Information digitisation has been carried out using the system

2 proposed by Bournaud et al. (1982), where 4 integer values are used to describe the degree of association between a species and the categories of a variable, for example, the categories of a habitat variable in the Macrohabitats file: 0 - no association, 1- minimal association (i.e. the habitat category is only marginally used by the species); 2- medium association (i.e. the habitat category is part of the normal range of the species); 3- maximal association (i.e. the habitat category is optimally preferred by the species).

The link between the spreadsheet files is provided by the species list, which is common to all of them, allowing sections of different spreadsheets to be joined together as required. A nomenclaturally correct list of the species covered by the database is also given, in Appendix 2 to the present volume..

Up to now, the need to regionalise the information about species coded into the spreadsheets has proved minimal. The most notable exception is coding of the flight period data in the Traits file. Variation in the length and timing of the flight period of many species, in different parts of Europe covered by the database, is sufficient to significantly reduce species predictability, were only a generalised flight period to be coded for each species. This has led to inclusion of sets of regional flight period codings for each species. Unfortunately, regional flight period data are not available for all parts of Europe covered by the database, so that generalised flight period coding still has to be used for some parts. It is anticipated that, as the coverage provided by the database expands, so will the need to progressively regionalise the coded habitat data. Already, certain habitat categories covered by the Macrohabitats file are only found in parts of the geographic area covered by the database. The most obvious examples are coastal habitat categories.

Although the main bulk of the present volume has been culled from material already published by the authors, it goes considerably beyond what has been published, particularly in its provision of background information.

So far, this introduction has been concerned primarily with the anatomy and coverage of the database, but it was felt that an attempt should be made to also present something of the philosophy behind the database, and the remainder of the Introduction is devoted to

3 that and allied issues. The database is essentially a tool for use in interpretation of data gathered in the field, so Chapter 2 of this volume is concerned with preferred field techniques employed for collecting adult syrphids, and their standardisation. It does not represent a review of all sampling methods currently in use. No attempt has been made to review procedures for sampling syrphid larvae in the field - although techniques are arguably available for use in a limited range of habitat/micro-habitat types, standardised larval collection methods are otherwise non-existent, or require substantial research effort to increase their reliability to an acceptable level. In Chapter 3 the processing of field- collected material is considered. Once again, this is not an attempt to review all available alternatives, but more an outline of a tried and trusted approach, which may be adopted by those wishing to deal with syrphid material collected using the techniques described in Chapter 2. Chapter 4 focuses on manipulation of the spreadsheets and a particular statistical treatment of results which is of potentially wide application in use of the database. Chapter 5 provides examples of use of the database in various contexts, demonstrating, in particular, what can be achieved without recourse to statistics beyond production of the humble histogram. The volume concludes with an overview of the database’s progress to-date, in Chapter 6. Three appendices to the volume are provided, in the form of Excel files. These are referred to at appropriate points in the main body of the text: Appendix 1 under section 2.1.1; Appendices 2 and 3 under section 3.2.

1.1. Invertebrates in environmental interpretation and evaluation.

The invertebrates play key roles in wetland ecosystem functions and processes (e.g. decomposition of organic matter, flower pollination, predation). They are also able to provide a holistic picture of the interaction between fundamental ecological processes and to rapidly adjust their occurrence and abundance following modification of their environment. Therefore, several invertebrate-based systems of bioevaluation have been developed. Examples for the aquatic environments are well known (Sladecek, 1973; Verneaux et al., 1982; Wright et al., 1984; Foeckler, 1991). Among terrestrial invertebrates, some groups, such as the carabid beetles, have been repeatedly used for site evaluation purposes Refseth, 1980; Luff, 1987; Eyre & Rushton, 1989; Stork, 1990). Furthermore, invertebrates have frequently been addressed in integrated studies of alluvial wetland systems, an example of which is the comprehensive study of the South Moravian floodplain forests in Czechoslovakia (Penka et al.; 1985, 1991).

4 Use of invertebrates in evaluation of terrestrial sites has been explored by various authors, using a variety of different approaches. Disney (1986) and Day (1987) focused on comparison between sites. Decleer (1990) and Brunel et al. (1990) concerned themselves with comparison between different parts of one site. Some authors (eg. Disney, 1986; Speight, 1986; Eyre et al., 1986) considered the relative suitability of different taxonomic groups for use in such studies. Siepel (1989) has sought to establish a method for assessing the efficiency of individual species as tools in site evaluation. More frequently, authors simply employ taxonomic groups known to them, without comment.

In Europe, the Syrphidae meet most of the criteria listed by Speight (1986), for selection of insect groups to use in site evaluation processes and additional criteria recognised more recently: a) Less than 5% of the genera pose significant identification problems and the taxonomic literature is readily accessible, although scattered. The species may be identified using external morphology. Following extensive revisionary work in the period 1960-80 and appearance of the relevant volume of the Catalogue of Palaearctic Diptera (Peck, 1988), a reliable nomenclature has emerged which is increasingly being used by European workers. b) Reliable, recent, species lists are available for various European countries, especially in western Europe and the entire European fauna has been catalogued recently (Peck, 1988), totaling approximately 700 species. There has been no pan-European study to establish to which IUCN status category each species should be consigned, but most recently-published national lists provide status data. c) Ecological information about the species is generally sufficient to characterise their habitat associations in terms of generally understood habitat categories and to demonstrate that the species exhibit a high degree of ecological fidelity. Syrphid faunas occur in nearly all terrestrial and freshwater habitats except cave systems, main channels of rivers and open waters of lakes. Larval microhabitat may be characterised for nearly all species. d) The range of generation times exhibited by different species (8 weeks to 2 years), coupled with their rapid mobility and various microhabitat preferences, results in the syrphid species on a site providing information about both short (e.g. seasonal) and longer term changes in site conditions. e) On-site sampling of Syrphidae can be standardised and carried out over short periods using commercially-available equipment. Storage of samples is undemanding in terms of

5 space, labour and facilities and processing of samples is rapid, such that complete results can be obtained within two months of a standard site visit.

1.2. The basic approach adopted in the syrphid database.

The keystone concept behind the data files and their structure is that enough is known of the habitat associations and other attributes of European Syrphidae for the syrphid fauna of a site to be predicted, from a knowledge of the habitats present on-site and the species recorded from the part of Europe in which the site is located. But the information coded into the database provides for a wide range of applications at site, landscape, regional, national and international levels. Such information is largely unused in traditional forms of treatment of species lists, in which the species names simply become integers in a largely statistical operation, divorced from all other information about the species themselves. In those circumstances, the questions which can be addressed are largely statistical rather than biological in nature and are circumscribed by the limitations of statistical techniques, rather than those of biological information, with analyses truncated by the difficulties of dealing with species represented by both large numbers of specimens or very few specimens, and numbers of specimens collected being regarded as of greater significance than the biology of the species they represent. The files in the syrphid database are structured to maximise the use of the biological information about the species, enabling the biological attributes of species collected to be compared and analysed, not just the relative frequency of the species. The attributes of predicted species can also be compared and analysed. The predictive process is equivalent to putting the European syrphid fauna through a series of sieves, with the species which pass through all of the sieves together constituting the final predicted species list. It is repeatedly referred to in different sections of this text.

1.3. The ingredients of the predictive process.

The predictive capabilities of the database can be used in various ways, only some of which will be explored in this volume. Its basic use is in conjunction with species lists derived from individual sites, such as protected areas or other areas in a natural/semi- natural condition. Interpretation of such species lists is liable to be of interest to a wide range of potential users of the database, from syrphid specialist to land manager and environmental consultant, and is judged to become one of the most frequent applications

6 of the database. So for introducing the predictive process, the approach to predicting a site species list is used here. In order to run a basic prediction of the syrphid fauna of a site, two sets of information are needed:

a) a reliable species list for the region within which the site is located, b) a list of the syrphid habitats occurring on the site.

Use of a predictive mechanism based on a species pool makes this approach akin to certain others, such as the English system used in running water assessment (Wright et al., 1984), or the "assembly rules" approach proposed by Keddy (1992).

1.3.1. Regional lists

A basic premise of the predictive process is that the fauna of a site is a sub-set of the fauna occurring in the “region” within which the site is located, i.e. that the site fauna is derived from the species pool of that region. The species pool relevant to a particular prediction process varies in its geographic coverage with the geographic scale at which the database is being used. Thus, for considering the significance of a site at European scale, the fauna of the entire land mass of Europe would be the appropriate species pool, while for considering its significance at national level the national species list would be the relevant species pool. For considering the management of a site within a National Park, the faunal list for the National Park might be the appropriate species pool.

During course of the FAEWE project regional species lists were needed for Ireland and central France. For Ireland, the list existed already and required only a small amount of updating (Speight & Nash, 1993, Maibach et al, 1994, Speight & Chandler, 1995, Speight, 1996a). For central France, the nomenclatural confusion surrounding the existing national list made production of a regional list impossible, without first establishing a verifiable list for the country as a whole. Revision of the French list took five years (Speight, 1993, 1994; Speight et al, 1998). The subsequently published regional list for central France (Speight, 1996b) comprised 223 species.Similar endeavours may be needed to produce reliable regional lists for other parts of Europe where the database is employed, although various lists are in existence already and may only require updating. A range of available national and other regional lists is incorporated into the database in the Range and Status volume.

7 1.3.2. Syrphid Habitats.

The array of syrphid habitats covered by the database is detailed in the Macrohabitats volume, which also discusses the difficulties of deriving generally applicable habitat categories. The habitat concept employed is essentially that its habitat is where a species can live out its entire life cycle. The term habitat is not employed simply as an expression of where the adults of a syrphid species can be found. Adult habitats in that sense are detailed in the Species Accounts volume. Each habitat category used is defined in the Macrohabitats volume, in the Glossary of Macrohabitat categories, which also shows the extent to which the categories recognised coincide with habitat categories recognised in the CORINE system ( and hence the EU Habitats Directive). The coverage is aimed primarily at so-called natural/semi-natural habitats and the database is least effective in landscape intensively used by man, because of the difficulties of identifying meaningful habitat categories. For instance, one man-made landscape feature which is easily recognisable is a quarry. However, if an attempt is made to code species according to the likelihood of their occurrence in quarries, it becomes immediately apparent that the condition of the quarry is more important than the fact that it is a quarry. A partially- flooded quarry will potentially support some species associated with temporary or permanent pools, while its sloping side-walls may support some species of dry grassland. It is more effective to classify the habitat representation on such totally artificial sites according to its most similar natural analogues, so that a quarry might be regarded as dry grassland with temporary or permanent pools. But even then, prediction of the associated species is not reliable, because man-made sites include combinations of features which do not occur naturally, as well as features which do not occur at all under natural conditions.

In order to obtain a knowledge of the syrphid habitats occurring on a site it is necessary to conduct a habitat survey. A habitat survey procedure is detailed later in this text, in the section relating to Field Procedures.

8 1.4. Modelling

Use of the syrphid database does not involved modelling as generally perceived. However, prediction of a site fauna depends upon a form of modelling of the site, in which salient site characteristics are observed and recorded in a standardised manner, using a classified system of habitat categories readily accessible to interpretation by the human eye and capable of differentiating a wide range of biotopes/ecosystems and their components from one another. These particular habitat categories have been selected also because syrphids respond to them i.e. they are features which may be used to describe differences between these species in their habitat requirements. Using the habitat data collected on-site in conjunction with the database thus involves a form of reconstruction of the site within the machine, linking each of the species in the database to that construction by means of the degree of association between the habitats and each species, as coded into the data files. This albeit crude model of the site is thus produced complete with its associated syrphid fauna, providing the prediction mechanism.

1.5. Origin and development of the syrphid database.

The syrphid database was developed during the STEP programme of the EU, under the project on Functional Analysis of European Wetland Ecosystems (FAEWE). Three taxonomic groups of invertebrates were employed as tools in the FAEWE project, Carabidae (Coleoptera), gastropod molluscs (excluding slugs of the families Milacidae, Limacidae, Agrolimacidae, Boettgerillidae and Arionidae) and Syrphidae (Diptera). Together, these groups comprised more than 700 species, in the geographic areas covered by the project (Ireland and central France). The method of their use was to transcribe biological and other information about them into databases and then design a procedure for interrogation of the databases in a prescribed fashion, in order to provide non- specialists with a mechanism for interpretation of invertebrate species lists, in particular species lists derived from sites located on river floodplains. For purposes of that project, database interrogation was progressively focused upon gaining an overview of a site’s condition, in terms of its degree of function in maintaining biodiversity as expressed by its invertebrate fauna, in so far as this may be adduced from the taxonomic groups covered.

9 In focusing upon assessment of site function in maintaining biodiversity, the immediate objective of the invertebrate studies of the FAEWE project was design and testing of a mechanism for integrating invertebrates into the functional analysis procedure being set up for the FAEWE project in general. This mechanism took the form of a “decision tree”, which allows a standardised form of interrogation of the database, and its use is shown later in this text.

At the end of the FAEWE project, the syrphid database was simply a set of Excel spreadsheets covering the syrphid faunas of Ireland and Central France, unusable as a referenced source of information and with an uncertain future. Since then, the coverage of the Excel files has been extended to the syrphid fauna of the entire Atlantic zone of Europe and beyond, a text file of species accounts of all the species covered has been prepared to accompany the spreadsheets and these files, together with explanatory material and the present text on use of the database, have together been published, so that the database may be cited by its users.

The 1999 version of the database covers the syrphid faunas and habitats of the Central and Atlantic Regions of the EU, and extends to provide partial coverage of the Mediterranean and Northern Regions. The material in the existing files is updated annually, to keep abreast of developments in our knowledge of the species, and to take account of criticisms and comments received. In an attempt to provide an automated outlet for dissemination of information about the database, a demonstration version was installed on an internet website shortly after termination of the FAEWE project. It was at this stage that the database acquired the name “Syrph the Net”.

10 Chapter 2. FIELD PROCEDURES

Field information of two types is used with the database: information about the character of the target site(s) and samples or inventories of the on-site syrphid fauna. In this context, a site may be defined as a piece of ground forming the object of a study or inquiry. As such, it is not necessarily a homogenous patch of a single habitat type recognised in the database, and field survey requires to be adapted to the degree of heterogeneity exhibited by the terrain under examination. Essentially, each habitat type represented requires both recording and sampling.

2.1. Site description.

This procedure is based on use of the Habitat Survey form provided in Appendix 1. Although it is possible to gain information on which syrphid habitats are present on a site, from surveys carried out by, for instance, botanists, the product is not usually satisfactory for use with the syrphid database, so that a habitat survey based on use of the Habitat Survey form is normally necessary, whatever other forms of habitat survey have been carried out on a site to serve the needs of other disciplines..

2.1.1. Use of the Habitat Survey Form

A Habitat Survey form was first provided as part of the 1998 version of the database. It has since been redesigned, because the previous version proved awkward to handle in the field and required a lot of space, if stored for reference purposes. The revised version requires a record to be made of the the habitats observed per sampling station, as previously, but recording the habitats by means of their code-numbers has reduced the size of each form to a single sheet. For easy reference to the code numbers for habitat categories, and to match habitats on-site with those in the database, it is necessary to take into the field not only copies of the Habitat Survey form, but also a print-out of the Summary Table of habitat categories used in the Macrohabitats spreadsheet (which lists the habitats and their code numbers). That Summary Table may be found in the Macrohabitat Associations text file. Use of the Habitat Survey form has also demonstrated that it is invaluable to have the Glossary of Macrohabitat Categories (also in the Macrohabitat Associations text file) available, while filling out the form in the field. This is particularly necessary because observers often have different interpretations

11 of habitats, but in order to use the database to maximum advantage it is necessary to use the interpretations laid out in the Glossary of Macrohabitat Categories.

The Habitat Survey form should be filled out in the field, while on-site. Experience shows that completion of the form for up to ten different locations on a site can be achieved in a day (once some familiarity has been gained with use of the form it may be completed for one sampling station within 10 minutes - the time taken to carry out a site habitat survey is dependent more upon the distance between sampling stations than on the time taken to fill out the form), and may be carried out at almost any time of the year that the site is not either flooded or covered in snow, though best results can be expected during the growing season for local vegetation. A complication is provided by temporary water bodies, which may add significantly to the diversity of a syrphid fauna, but which, by definition, are only observable at certain times of the year. Ideally, habitat survey would be carried out twice on a site, once during the period of annual high water-level for ground-water and then again during time of low ground-water level. Failure to recognise the presence of temporary water bodies (seasonal streams, springs, flushes and pools) can lead to significant under-prediction of a site fauna and consequent failure to recognise the potential for some of the species recorded from a site to actually breed there.

Completion of Habitat Survey forms well in advance of any sampling programme can be valuable, in that it allows prediction of the most appropriate periods of the year in which sampling programmes might be conducted, from the information on seasonal availability of the predicted fauna provided in the Flight Period tables in the Traits spreadsheet.

Some general habitat categories are un-necessary to record in the field, because their presence can be adduced from the presence of other recorded categories. For instance, if category 11211 (mesophilous Fagus forest) is recorded for a site, on a Habitat Survey Form, this automatically means that the more general categories 1121 (Fagus forest), 112 (mesophilous/humid deciduous forest), 11 (deciduous forest) and 1 (forest) can be recorded from the site, so these categories do not require to be separately recorded on the form.

On the Habitat Survey Form, the categories to be recorded are of two types, macrohabitats and supplementary habitats, as in the Macrohabitats file. The form allows

12 recording of each supplementary habitat found in association with a macrohabitat. It is necessary that the supplementary habitats associated with each macrohabitat are recorded, since these associations have a significant influence on the potential constitution of a site fauna. For instance, a brookside in grassland can have a very different associated syrphid fauna from a brookside under the canopy of a forest, and a brook may pass from within a forest out into grassland within one site, or be present on a site in association with one of those macrohabitats but not the other.

In order to record the data collected on Habitat Survey Forms it is advisable to set up a separate Excel file of the Macrohabitat categories in the Macrohabitats file, into which the site survey data can be transcribed.

2.2. Field-sampling procedures

In the case of Syrphidae, field-sampling is dependent upon collection of the flighted adults. For operating the database, sampling procedure has been standardised around use of the Malaise trap as the sample unit. The relative efficiency of various trap designs in the capture of different sorts of flying insect is reviewed by Southwood (1978) and Muirhead-Thomson (1991).These authors do not specifically consider trap efficiency in relation to capture of Syrphidae, but they do demonstrate that any trapping mechanism has its own bias and that each taxonomic group responds somewhat differently to any particular trapping technique and regime. In deciding upon the Malaise trap as the standard sampling unit to use for Syrphidae, the following points were taken into consideration: a) analysis of results is dependent upon adequate samples of the local fauna, not a complete inventory of the local fauna, b) The analysis procedures employed depend primarily upon use of presence/absence data, c) Ease of transport of trapping equipment to and from possibly remote sites, rapidity of installation and removal of trapping equipment and ease of servicing of equipment installed are all of primary concern, d) Rapidity and simplicity of sample handling is important.

Little information about use of Malaise traps is yet available in the literature.The standard, commercially available Malaise trap can be erected and maintained by non-

13 specialists, with very little prior training. Similarly, on-site servicing of the traps and sample collection can be carried out swiftly and simply by non-specialists. Further, the plastic bottles attached to the traps, and into which samples are collected, can be used for transport of the samples and for storage throughout the sample processing phase. While installed on a Malaise trap, a collection bottle is part-filled with 70% alcohol or some similar preservative, into which the collected specimens fall.This also provides for preservation of the sample during transport from the field and subsequent laboratory processing. There is thus no need for transfer of samples from one container to another, from the time the bottle is installed on the trap until it is in use in the laboratory. Collection of samples and their transport can be carried out by non-specialist personnel.

While there is no standardised methodology for use of Malaise traps in sampling syrphid faunas, various applications of Malaise trap survey to work on Syrphidae are exemplified in the literature, from the scale of national distribution survey (e.g. Verlinden and Decleer, 1987) to site investigations (e.g. Haslett, 1988). Other authors have used water traps (e.g. Chemini et al, 1983), or hand nets (e.g. Kassebeer, 1993; Marcos-Garcia, 1990), but in these cases longitudinal surveys have been undertaken, carried out intermittently over months or even years. On adequately protected sites where agreement with land owners has been reached as to when and where a Malaise trap survey will take place, the Malaise trap provides rapid results without requiring constant on-site presence of a specialist (as required for hand-net survey) or frequent collection of samples and trap maintenance (as is required for water-trap survey). On inadequately protected sites, Malaise traps, being highly visible structures in most landscapes, are highly susceptible to vandalism or removal and to damage by livestock.

2.2.1 Placing of Malaise traps.

Where and whether insects fly is determined by many factors, including climate, time of day and site topography. When in flight they are not evenly distributed, within that fraction of the air column above a site intruded upon by flight interception traps like the Malaise trap. The position in which a Malaise trap is installed and its orientation thus influence its efficiency. Basically, Malaise traps can be positioned either on or off flight lines and either orientated or not along a north/south axis. Flight lines are largely dictated by local micro-topography and the location and direction of many of them can be detected by human eye, from juxtaposition of site features. It is evident from the work of

14 authors such as Aubert et al (1976) and Gatter and Schmid (1990) that positioning a Malaise trap across a flight line maximises the catch of syrphids flying through a site from elsewhere (including migrators). Conversely, placing a trap off flight lines maximises the catch of syrphids engaged in local, on-site movements. Orienting a trap north/south, with its high point facing south, maximises catches of insects liable to fly towards the point of highest light intensity (i.e. the sun) on contact with a trap (i.e. heliophile, day-flying insects like syrphids). Trap alignment in a north/south direction can be achieved using a compass.

Site factors operating over short periods can also have a significant effect on Malaise trap catches, for instance a large patch of some low-growing plant which comes into bloom in the vicinity of a trap during a trapping campaign can greatly increase the number of syrphids caught. Conversely , the efficiency of Malaise traps left in situ for an entire flight season (i.e. spring to autumn) can be reduced by change in the condition of the ground vegetation as the growing season progresses. Installation of traps in a crop of maize (Zea mais) provides an extreme example - at the beginning of the season the ground around the trap is virtually bare, whereas at the end the maize is higher than the trap itself, having clear implications to its accessibility as an interception trap for flying insects. In deciding where to position a Malaise trap, such features can be either sought or avoided, as a matter of choice, but cannot be simply ignored.

In conducting a short duration (e.g. ten-day) field campaign, it might be considered self- evident that Malaise traps should be positioned to obtain the maximum quantity of data in the minimum time. However, as indicated in the previous paragraph, maximising the trap catch is not necessarily synonymous with maximising the catch of species which have developed locally, for instance, and the questions to be answered by conducting the trapping programme require to be considered carefully in deciding trap placement. In order to overcome the potential influence of trap placement on trap catch it is advisable to use Malaise traps in pairs, the two traps of a pair being placed close to each other (though sufficiently far apart that they do not interfere with each other’s action) at the chosen trapping station. The degree of similarity between the catches made by two traps installed at a trapping station can be ascertained, and compared with the catches of traps from other trapping stations, to verify that the catch of a particular trap is less affected by its placement than by the character of its surrounding habitats.

15 The need to sample the fauna of each principal habitat type observed on-site determines the minimum number of Malaise traps to be positioned there - it being advisable to place at least one pair of traps within the area occupied by each of the observed habitats. It may be necessary to install traps at additional locations, as required by co-workers.

2.2.2 Sample collection and trap maintenance.

Sample collection from a Malaise trap entails simply unscrewing and capping the collection bottle and transporting it to the laboratory. In temperate conditions 70% alcohol makes an acceptable preservative for use in collection bottles, but in the warmer conditions of the summer months of central France a 30% solution of the less volatile ethylene glycol is more appropriate. Windy conditions can also result in higher evaporation rates of alcohol and can make ethylene glycol a preferrable option for use in the field, in Malaise trap bottles. If ethylene glycol has been used, it is desirable to strain it from the caught insects and replace it with 70% alcohol within 3 weeks from the date at which the collection bottle was put in place on the trap, to prevent disintegration of specimens.

It is advisable to check collection bottles in place on traps at least once every two weeks, and once a week in conditions of high wind or high temperature, in case there has been increased evaporation of preservative. Catch rate varies considerably and there is also need to ensure that bottles do not fill with collected insects to above the surface of the preservative. In conditions where rapid catch rates might be anticipated it can be necessary to check bottles every few days. In exceptional circumstances it may be necessary to replace bottles on a daily basis. Correct labelling of collection bottles is critically important, to ensure it is known from which trap each is derived. To help ensure that the sample bottle collected from a trap is labelled correctly, it is advisable to mark the permanently attached upper bottle on the Malaise trap with the code name for that trap, using non-water-soluble ink. This code name is then immediately available for reference when the sample-bottle is labelled, which should be undertaken either as part of the process of attaching the sample-bottle to the trap, or as part of the removal process, to ensure there is no confusion between sample-bottles from different traps. It is preferable to use code systems which can be used to refer to particular traps, or their products, throughout the field and laboratory procedures, so that the code name attached to a trap

16 may finally be used for the column(s) referring to the syrphids recorded from that trap, in the Excel file set up to hold the transcribed field data in the computer.

In most instances damage to traps is limited to guy ropes being severed or pulled out of the ground. This rarely results in loss of a sample, but can reduce trap efficiency. If traps carry waterproof and sunlight-stable notices, explaining their purpose and asking for co- operation, incidence of vandalism is surprisingly rare. But when it occurs it almost invariably results in loss of samples. High wind can also wreak havoc and any trap which suffers from the attentions of one of the larger forms of domestic stock, such as cows or horses, can be totally destroyed. Traps can only be effective in the presence of these large animals if protected from them by strong, temporary fencing or electric fencing - or by reaching agreement with landowners which results in domestic stock being grazed elsewhere for the duration of a sampling campaign.This latter alternative it eminently preferable to erection of temporary fencing of any sort, which tends to be both very time- consuming and unreliable. For any field campaign it is advisable to hold a few spare traps in reserve, in order to guard against possible trap destruction. Under most circumstances, the time taken to complete a trap round is largely dependent upon the distance between traps and how closely they may be approached by vehicle, rather than the time required to service the traps themselves.

2.2.3 Field Sampling Strategy: obtaining representative samples of site faunas

There are various factors that require to be considered in designing a fieldwork campaign aimed at obtaining a representative sample of the syrphid fauna of a site, using Malaise traps. Survey aimed at inventorising the syrphid fauna of a site requires less rigorous consideration of optimal sampling periods, but is more demanding of man-power and time.

2.2.3.1 Choice of sampling period

Over most of Europe, adult syrphids are on the wing between April and September (inclusive), so Malaise-trap sampling outside this period is not practical, except within the Mediterranean zone. A field campaign which had to be conducted during the winter could not usefully include adult syrphids in site investigation processes. Within the period April-September the various species are in flight at different times, and unless

17 sampling can be carried out throughout that period, choice has to be made of when to sample. Most univoltine species are only in flight at the beginning of the summer, whereas polyvoltine species recur again later in the year. This is illustrated in Figs. 2.1 and 2.2.. So, to sample univoltine species the optimal period is April/beginning June, over most of Europe. Figs 2.1. and 2.2. also indicate that a second sampling campaign, in July/August, might be expected to show which polyvoltine species move into a site during the summer, though absent there earlier in the year.The periods of the year optimal for sampling can also be influenced by the type of habitat in which sampling is being carried out. This is illustrated for the potential fauna of the FAEWE site at Decize in Figs. 2.3. and 2.4..

In Fig.2.3, the flight season data for all the species represented on the regional list for central France that are associated with habitats observed on the Decize site have been put together, to give a composite flight season profile. This shows that, in the latter half of May/first half of June, the number of species available there should be at a maximum, suggesting this would be the optimal period for a field campaign.This composite flight profile is typical for most habitats in atlantic parts of Europe, so in principle it is true that in this region of the continent the most opportune time for sampling a syrphid fauna is end May/beginning June. However, this can be a period of very variable weather and in years in which spring is retarded the fauna is as well, reducing sample catches. Under such conditions, sampling should be postponed to mid-June, if possible.

In Fig. 2.4. two groups of species contributing to Fig. 2.3. have been taken separately: the flight season profile of the species associated with the mature/overmature alluvial softwood forest has been compared with that of the unimproved pasture species. Fig.2.4. shows that while the period end May/beginning June is an optimal sampling period for species associated with both types of habitat on the Decize site, the pasture species might also reasonably be sampled through into July/August, whereas the forest species would be only half as well represented during this period as in end May/beginning June. In order to optimise a sampling programme it is necessary to have a clear understanding of which habitats in the target area are the particular objects of concern. In the event that free choice of sampling period cannot be exercised, it is necessary to take into account the flight period profile of the fauna associated with each habitat observed on site when analysing results. This can be achieved using the database.

18 2.2.3.2 Duration of field campaigns

Optimisation of field campaign duration brings into consideration such issues as the cost of a field campaign and the length of time a farmer, or other land owner/user, might reasonably be expected to modify his/her schedule for use of a site in order to accomodate survey needs. These logistical considerations dictate that on most sites no Malaise-trap-based on-site sampling campaign should continue for more than a few weeks. In a nature reserve, national park or other protected site, field campaigns can reasonably continue for much longer than this, but only a proportion of sites requiring investigation are likely to fall into these categories. Working on the basis that installing an average set (twenty) of Malaise traps takes two days and that a similar length of time is also required to remove them, based on expert advice fourteen days has been identified as the minimum period in which a Malaise trap field campaign could usefully be carried out. This then provides for a ten-day sampling period, which allows for the occurrence of flight-inhibiting weather (e.g. high wind or rain) for part of the period. Two such field campaigns, carried out within the period beginning June/end August and giving together a total of 20 days of sample collection, have similarly been taken as the minimum required to amass an adequate sample of the syrphid fauna of a target site (see below). In most circumstances the optimal timing of these two sample periods is probably June and August.

Fig.2.5. shows how the number of species collected increased as the sampling period was extended, during course of a longitudinal Malaise trap survey conducted in an Irish National Park. As might be expected, with the addition of each ten-day sampling period the total number of species collected continued to rise, from June through to September. However, after the first 20 days of sampling more than half the total number of species had been collected and from then on the species increment resulting from each additional ten-day period reduced. Fig.2.6. shows what would have been the effect of timing the first of two ten-day field campaigns in the second half of June and the second field campaign at each of the subsequent ten-day periods through to September. Whatever dates the second ten-day sampling period covered, the number of different species collected by the two campaigns put together amounts to at least half the total number of species trapped in the entire season June/October.

19 Longitudinal sets of Malaise trap-collected data like those derived from the Killarney Park are not available from many sites, so it is uncertain how typical the Killarney results may be. However, they do show that 20 days' sampling by Malaise trap within the period mid June/September can collect half the species available to the traps during the entire 110 day period mid-June/October. Precht and Cölln (1996a,b) report on a longitudinal Malaise-trapping exercise conducted by themselves on a site in Germany, with more-or- less the same objective. They conclude that 80% of the syrphids which may be trapped on a site using Malaise traps may be collected by four weeks of Malaise trap activity in June/July. A similar data-set from one site in New Zealand was used by Hutcheson (1990), as a basis for concluding that 28 days of Malaise-trapping represent an adequate basis for sampling flying beetles (Coleoptera) for purposes of site characterisation.

2.2.4. Inventory survey

Survey carried out, with the objective of compiling an inventory of the syrphid fauna of a site, requires to accommodate the requirement for all habitats (including supplementary habitats) present on the site to be sampled and through the flight period of all the species predicted to occur on the site. The array of habitats present can be established by habitat survey carried out in advance of installation of the Malaise traps. In inventory work it is more important to have the Malaise traps in place before, or at least by, the earliest date predicted for the onset of the flight season, because there can be as much as a month difference between years in the date at which the early spring species actually start to fly. Installation of traps by some later date, at which most of the early species would be supposedly on the wing, could easily result in a number of them being missed, in a year in which spring started earlier than usual. This is particularly true, given that so many of the early spring species are also univoltine, and so cannot be collected at any other time of the year.

Even though it is advisable to obtain samples from all parts of the flight season, when understaking inventory work, this does not imply a need for continuous sampling. In general, one 20-day time unit of sampling within each month of the flight season should prove adequate. It is advisable to use 20-day time units, rather than 10-day units, to minimize the frequency with which it is necessary to visit the traps.It is to be remembered that, if an inventory survey results in collection of a superabundance of material, it is always possible to process only a subset of the collected material, if that is

20 deemed adequate for compilation of the inventory. But, if insufficient material is collected, there is no easy way to deal with the situation other than to carry out supplementary survey work, an option which is frequently unavailable.

Malaise traps are not mobile and habitat survey is fallible, so that it is well possible for certain habitats on a site to be too distant from any Malaise trap emplacement to ensure their fauna will be comprehensively collected by the traps. To minimize such effects it is helpful to employ a second collection method, when conducting inventory work by means of Malaise trap survey. Collection by direct observation, using an insect net, carried out by an experienced syrphid worker, provides the ideal supplement, since the experienced observer both can and will visit parts of the site remote from the traps. If the Malaise traps are operating efficiently, supplementing their catch by insect net work is unlikely to add dramatically to the resultant species lists. But if there are elements of site heterogeneity that have been inadvertently overlooked in placing the Malaise traps, or which it has been impractical to sample using Malaise traps, then collecting by net should help to ensure that any part of the syrphid fauna of the site that is dependent upon such neglected site elements will be added to the inventory.

90 80 70 60 50 40 30 20 10 0

Fig.2.1.: Relation between flight period and number of generations/annum: central France species pool, showing number of species in each generation category on the wing in each month. Rectangles denote species with no more than 1 generation/annum; diamonds denote species which may be univoltine or divoltine; triangles denote species with two or more generations/annum.

21 70

60

50

40

30

20

10

0

Fig.2.2.: Relation between flight period and number of generations/annum: Irish species pool, showing number of species in each generation category on the wing in each month. Rectangles denote species with no more than 1 generation/annum; Diamonds denote species which may be univoltine or divoltine; Triangles denote species with two or more generations/annum.

250

200

150

100

50

0

Fig.2.3.: Combined flight period profile for all syrphid species associated with the habitats observed on the FAEWE Decize site and represented in the central France species pool. Derived from the flight season data-file for central France, used with the species list for central France and the list of syrphid habitats observed on the Decize site.

22

120

100

80

60

40

20

0

Fig 2.4.: Flight period profiles of the syrphid species associated with two different habitat types observed on the FAEWE Decize site compared. Hollow columns = unimproved pasture species; striped columns = mature/overmature alluvial softwood species.

45 40 35 30 25 20 15 10 5 0

Fig.2.5.: Showing increment in number of species collected with increased length of sampling period, using Malaise-trap data from the Killarney National Park, Ireland. Each contiguous sampling period is ten days. During the first sampling period, 16-26 June 1993, 15 species were collected. Sampling continued for 110 days and by the end of the last sample period a total of 42 species had been collected.

23

35

30

25

20

15

10

5

0

Fig.2.6.: Total number of different syrphid species obtained from combining the results of a field campaign 16-26 June (first column) with those of a second field campaign of the same length at different dates up to 7 October, using Malaise trap data from the Killarney National Park, Ireland, in 1993. The total number of species collected 16 June-4 October was 42. 1 = 16-26.6; 2 = 26.6-6.7; 3 = 6-16.7; 4 = 16-26.7; 5 = 26.7-5.8; 6 = 5.8-15.8; 7 = 15-25.8; 8 = 25.8-4.9; 9 = 4-14.9; 10 = 14-24.9; 11 = 24.9-4.10.

24 Chapter 3. LABORATORY PROCEDURES

Although the actual techniques employed in the laboratory depend on which taxonomic group is involved, the same processes are common to all groups. The target organisms have to be sorted from among other invertebrates and debris collected with them, stored, determined and recorded for transcription into the computer.

3.1. Treatment of samples

A collection bottle removed from a Malaise trap can be used to store the insects collected until the samples are required for sorting. Sorting is best carried out under a binocular microscope, using a petri dish to sort, one after the other, decanted fractions of the material from the collection bottle, until the entire content of the bottle has been examined. The alcohol in the collection bottle is likely to become clouded by shed scales from the wings of Lepidoptera collected and it is advisable to strain this off through a fine sieve and sort using clean alcohol in the petri dish. If the alcohol strained from the collection bottle is filtered it may be reused for storage of collected material. Clear and unambiguous labelling of each container storing collected material is vital - confusion between material collected by different traps, or at different dates, renders samples unusable. The syrphid specimens extracted from a sample are best retained (in alcohol) in a separate, smaller, closed container, labelled to show their sample of origin, so that they may be determined together with the syrphid material extracted from other trap samples from the same field campaign. Following determination, the syrphid sub-sample sorted from a collection bottle may be re-united with the parent sample, but it is prudent simply to insert the small, closed container holding the syrphids into the parent sample bottle as a discrete entity, in case there is need to re-examine one or more of the syrphid specimens at a later date.

Sorting syrphids from other insects collected by the traps is time-consuming and demands a trained eye. An experienced sorter can be expected to sort syrphids from up to 300ml of insect material in four hours, and 10 collection bottles can take from 2 days to a week to sort.

25 3.2. Determination of the sorted specimens.

For use with the database, specimens have to be identified accurately and to species, thus requiring specialist attention. The need for accurate determinations is paramount, since use of the procedure is entirely dependent upon species-related information. The process of determination is time-consuming. With experience, the material extracted from 10 collection bottles all derived from the same local fauna can usually be determined to species in 1 or 2 days. But extra time may be needed for particular specimens. In some instances, there may be need to remove specimens from alcohol and dry them, prior to determination, because the existing identification keys are all based on dry specimens. For an inexperienced observer, the process of determining syrphids preserved in alcohol is a frustrating, tedious business, fraught with difficulty. There is no recent publication in which all known European genera of Syrphidae are keyed out. However, there are quasi- regional works which cover most genera. The most comprehensive accounts are those of the following authors: Bradescu (1991), Stubbs & Falk (1983), Torp (1994), van der Goot (1981), Verlinden (1991), Violovitsch (1986), Vockeroth & Thompson (1987). Which genera are covered by each of these accounts is detailed in tabular form in Appendix 3 to this volume. In addition to indicating which of the genera covered by the database are keyed out in those publications, Appendix 3 also shows which other European genera they key out. For completeness, certain genera whose range is peripheral to Europe are included in Appendix 3. These genera comprise Allograpta, Asarcina, Ischiodon and Megaspis. Further information on identification is given in the Species Accounts volume, under the sub-heading “Determination” included in the account of each species. That focuses, in particular, on literature for determination of individual species. The Species Accounts also provide details of literature available for identification of developmental stages, under the subheading “Larva”.The most comprehensive recent texts dealing with identification of larvae are those of Rotheray (1994, 1999) and Torp (1994).

For purposes of publication, the names of species listed as occurring in a site or region normally require to be quoted complete, i.e. followed by both name of author and date of description. Putting this information together for even a small number of species can be an irksome task, so a complete list of nomenclaturally correct names of the species covered by the database is incorporated into Appendix 2 of this volume. The species

26 listed there are the same, and in the same order, as in the other Excel files in the database, so that the list in Appendix 2 may be used in conjunction with the other spreadsheets.

Papers on taxonomy and ecology of European syrphids are scattered through a vast range of scientific periodicals. Only the recently-established journal Volucella deals exclusively with the syrphid fauna of the western Palaearctic. Volucella provides an outlet for autecological, distributional and taxonomic information on the European species and an increasing proportion of material for up-dating the syrphid database is now coming from its pages. Further information about Volucella may be obtained from: Dr.U.Schmid, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany.

3.3. Recording the determined specimens.

It is advisable to retain the most detailed record possible of the species identified in each sample, and in such a way that transcription of the recorded information, into an Excel file which may be used with the database, can be carried out simply.

Part of a simple recording form is shown in Fig.3.1 The format used allows the syrphid specimens from each Malaise trap sample to be recorded on a separate record sheet. It is imperative that the trap sample from which the syrphid records are derived is also indicated on each record sheet, preferably using some code which can employed to denote this trap sample in the Excel file to which the recorded syrphid data are to be transcribed.

TAXONOMIC GROUP SITE Trap No.: DATE

Species no.males no.femal remarks es

27 Fig.3.1: Part of a form for recording syrphid specimens from a trap sample. Chapter 4. DATA PROCESSING PROCEDURES

The spreadsheets are set up for use with species lists, species lists being regarded as the most likely type of information generally available about the syrphids of a site or region. This does not preclude the use of the spreadsheets with quantitative data. But interpretation of quantitative data may be treated as a separate issue and the logic implicit in the approach adopted here is that once species presence/absence data can be interpreted, then it becomes worthwhile to consider quantitative information, with all the additional problems of interpretation that entails. The basic procedure outlined here assumes use of presence/absence data.

4.1. Field data files

The need to transcribe site data into spreadsheets has been alluded to earlier in this text. If the data on habitats observed on-site are transcribed to an Excel file, with the habitat categories arranged exactly as in the Macrohabitats file, this can facilitate extraction of the habitat array required for carrying out the prediction procedure for a site.

The species-observed data for sites can all be transferred from record forms to one Excel file set up to hold the information. The observed-species categories should be clearly labelled to avoid confusion between them. If the species are listed as in the other Excel files in the database, this greatly simplifies file manipulation procedures..

4.2. The basic site interpretation procedure

As indicated earlier in this text, for purposes of employing the database a site may be defined as a piece of ground forming the object of a study or inquiry. Parts of sites from which data are collected by some sampling procedure are then considered as sub-sites or sampling stations. Interpretation of species lists from sub-sites, sites, groups of sites or regions can be carried out using the same basic procedure. The flow diagram in Figure 4.1a illustrates the first steps in a basic site interpretation procedure.

28 a) the species associated with the habitats observed on site are extracted from the regional list; using the habitat coding to extract only expected species i.e. those coded “2” or “3” for the relevant habitat categories, b) the species potentially in flight at the time of the field survey are extracted from the list obtained in a), using the relevant flight period categories in the Traits file. This provides a list of predicted species for a given site and a given sampling period c) the list of species actually sampled during the field survey is then compared with the

list of predicted species obtained in b).

REGIONAL LIST OF

SPECIES

Habitat Flight preferences of period data

the species

HABITAT List of predicted





























TYPES List of predicted species for a given

OCCURING species sampling period

ON SITE

LIST OF OBSERVED

SPECIESFOR A COMPARISON












































GIVEN DATE(field

data)

Figure 4.1a. Basic site interpretation procedure: production of the list of predicted species and its comparison with the list of observed species..

The comparison can be conducted using a great many different combinations of data categories from the spreadsheet files of the database. The actual combinations of categories used in any one instance will depend upon the requirements of the investigation. Fig.4.1b shows diagramatically an example of the procedure involved, which is very reminiscent of employing a catenary series of sieves of decreasing mesh- size, with data categories from the spreadsheet files taking the place of the sieves.

29 The example used in Fig.4.1b assumes the investigation relates to a fen site in Ireland, for which a reliable inventory of the syrphid fauna exists. The first operation is to make up a spreadsheet work file containing the relevant data categories, which in this instance are taken to comprise the following: 1. A copy of the database species list which accompanies the spreadsheet files, 2. The list of species observed on the site (from the file set up to hold site data), 3. The “Ireland” category, “European Range” variable, Range and Status file. 4. The “Fen (gen.)” category, “Wetlands” variable, Macrohabitats file. 5. A copy of the Microsite Features file (entire). 6. A copy of the Traits file (entire). 7. The 3 categories from the “Status, Ireland” variable, Range and Status file.

Sorting the work file, using the Irish species list category (sort 1), reduces the number of species considered to only those which occur in Ireland. Sorting the work file again, using first the “fen” category (sort 2), further reduces the number of species to be considered, to those which might be expected in fen in Ireland. This then is the predicted site-species-list, since on this imaginary site there is only the one habitat “fen” represented. By the same sorting process, the list of species observed on-site has been reduced to those associated with fen, so the list of on-site species associated with fen may now be compared with the list of species predicted for this habitat. The example assumes that, from direct comparison between these two lists, it is decided to examine certain differences between them further, commencing with a comparison between the expected and observed species associated with the “on tall herbs” category from the Microsite Features file. Sorting the work file using this category (sort 3) reduces the observed and predicted lists to species associated with tall herbs in fen, so that this comparison may be made. Traits of these species may then be considered and, in this instance, it is decided to compare the representation of species with predatory larvae, in the predicted and observed lists, using the “living animals” category from the “Food source (larvae)” variable (sort 4). Finally it is decided to consider the representation of threatened species among the predicted and observed species with predatory larvae on tall herbs in fen (sort 5). At this final level, the number of species involved may be very small.

This example is of a simplified case, in that it would be rare for only a single habitat category to be represented on a site, or for only one microsite feature or trait category to be brought into consideration during the process of comparison. For instance, it would be

30 more normal to consider the representation of all three categories of larval feeding type (predators, plant feeders and detritivores) at once. A more complex procedure then results, producing a dendritic structure to the comparison process and ending with a mutliplicity of final products for comparison.

It is necessary to take into consideration the differences between lists of observed species based on production of inventories, and lists of observed species based on production of representative samples, in conducting a comparison between predicted and observed lists. The types of comparison which can be meaningfully conducted are fewer when representative samples are involved, than when inventory lists are involved.

Atlantic zone species + habitats + traits + microhabitats + status + site species list c400 species Sort 1: regional list- Ireland

↓↓↓↓↓

Irish species + habitats + traits + microhabitats + status + site species list 171 species Sort 2: habitats on-site - fen ↓↓↓↓↓

Irish species predicted in fen + traits + microhabitats + status + species from fen observed on site Sort 3: Microhabitat -on tall herbs ↓↓↓↓↓

Irish species predicted in fen, with larvae on tall herbs + traits + status + species from tall herbs in fen observed on site Sort 4: Trait - predatory larvae ↓↓↓↓↓

Irish species with predatory larvae living on tall herbs, predicted in fen + status + species with predatory larvae on tall herbs in fen observed on site Sort 5: Status, - Ireland, threatened ↓↓↓↓↓ Threatened Irish species with predatory larvae living on tall herbs, predicted in fen, plus threatened species with predatory larvae on tall herbs in fen, observed on site

Fig.4.1b: Progressive sorting procedure, used in comparing species lists (diagrammatic).

4.3. Statistical/analytical techniques

31 No advanced statistical techniques are needed in order to carry out most of the procedures presented here, including the predictive procedure which is intended as a general use of the data base - the prediction of a site fauna and its comparison with the species actually sampled are solely based upon counts of numbers of species, and percentage calculations. However, the data files can be explored and described with appropriate multivariate ordination techniques, and coupled with the results of site surveys following the procedures set up by Dolédec & Chessel (1994) or Chevenet et al. (1994). An example of such multivariate investigations using the ADE software is given below, derived from Castella & Speight (1996). The ADE software is available free of charge, and may be downloaded from the following internet address (URL): http://biomserv.univ-lyon1.fr/ADE-4.html

4.3.1. Use of multivariate ordination techniques with the database

In this example, syrphid microhabitat and biological trait data are incorporated into the analysis of field data, providing insights into the extent to which particular species attributes are manifest in the fauna of different sampling stations. All the calculations were carried out using the ADE software. The procedures described follow the guidelines suggested by Dolédec and Chessel (1993a and b) and Chevenet et al (1994).

The first step is ordination of each of the two input data sets: - field data tabulating occurrences (as presence or absence) of the 78 syrphid species recorded at the 13 stations sampled during the two field programmes carried out on the French FAEWE sites, - microhabitat and trait data (78 species x 6 variables totaling 28 categories), extracted from the microhabitats and traits spreadsheets. For the purpose of the present example only a restricted number of categories of microhabitat and traits data are included in the analysis.

Characteristics of the French FAEWE sites and the sampling stations used there may be summarised as follows:

Four sampling stations were selected on each site on the basis of geomorphological, pedological and hydrological characteristics. These "HGMUs" (stations L1 to L4 and A1 to A4 for the Loire and Allier rivers respectively) were located along a 200 metre long transect, perpendicular to the river, starting at its bank and ending on the higher terrace of the floodplain. Each station was equipped for the regular

32 measurement of physico-chemical characteristics of the soil and groundwater and served as a reference point for the various experiments carried out within the frame of the project. The macroinvertebrate fauna was sampled at these 8 stations and at 5 supplementary stations (i.e. not installed for soil measurements): L0 and L2a on the Loire transect, AI, A0 and A5 on the Allier transect. These various sampling stations can be briefly characterised as follows: AI: riparian soft wood forest located on an island of the R.Allier L0 and A0: muddy sand (L0) or sand (A0) bank on the shore of the river, partly shaded by the adjacent Phalaris (L0) and scrub Salix (A0, L0). These stations were influenced by the frequent and rapid oscillations of the river water level and annually flooded. L1: Phalaris arundinacea and Salix sp. scrub on a sandy mud; partly shaded and annually flooded, being with LO on the lowest part of the transect. L2, A1, A2: riparian soft wood forest (Populus and Salix) with Urtica dioica (L2, A2) or Phalaris arundinacea (A1), flooded during spring floods. L2a: on the shore of an abandoned channel of the R.Loire (upstream disconnected, but almost permanently connected downstream with the river) and therefore as frequently flooded as LO or L1. Shaded by the riparian forest described for L2. L3: an extensively and not permanently grazed pasture on the higher alluvial terrace with a well drained sandy soil. Flooded only occasionally and submerged for only very short periods (1 to 2 days). A3: a more intensively grazed and humid pasture. Flooded each year (up to 10 days a year). L4: a depression (2 to 3 m deep) located in the pasture L3. The substrate was sand, locally covered with a shallow, poorly drained soil. This station was vegetated with grass and could be overflooded for periods of several days or weeks each year by rise in groundwater level. A4: a shallow depression with sedge communities, in the pasture A3 . A5: on the shore of a former channel of the Allier (oxbow lake with hydrophyte communities).

4.3.1.1. Reciprocal ordination of the species and sampling stations

The field data are processed by Correspondence Analysis (CA) Greenacre, 1984), which provides a reciprocal ordination of the species and of the sampling units. Correspondence Analysis was chosen instead of its detrended version (DCA) for several reasons, but in particular because: i) control over the geometry is lost in the process of detrending and DCA breaks up the optimal properties of CA, such as the maximisation of correlations or the reciprocal averaging (Greenacre, 1984, Lebreton and Yoccoz, 1987); ii) DCA is a rather arbitrary adjustment of CA and lacks mathematical coherency (Wartenberg et al., 1987; Digby and Kempton, 1987), hence it does not allow subsequent inter-battery analyses.

Fig..4.2 shows that, in this example the first two axes from the Correspondence Analysis ordinate the 13 sampling stations along a diagonal line from dense, shaded softwood riparian forest (A1, A2) at the top right to more open sites like dry unimproved pasture (L3, L4) or muddy sand flat along the river (L0) at the bottom left. This gradient can be associated along F1 with the increased occurrence of forest-dwelling species like Syrphus vitripennis or Myathropa florea and the parallel decrease of the open-ground / grassland 33 dwellers (e.g. Sphaerophoria scripta). The poorly-drained pasture A3 is an outlier to this gradient, with a particular assemblage mixing open ground species like Chrysotoxum arcuatum or Eumerus tuberculatus, together with forest dwellers, like Temnostoma bombylans, Brachyopa scutellaris or Meligramma cincta, the adults of which visit the pasture to feed upon flowers. Another reading of this ordination plane evidences for both sites (Allier and Loire) the repetition of the same ordination pattern for the four major sampling units from right to left along F1 (thick lines in Fig. 4.2) and a tendency to segregate these two transects along F2.

A3 F2

A1

A2

AI

F1

L1 A5 L2 AO L3 A4

LO L2a

F1

0.70- F2

Eigen

1.6

values

-1 1.8






















L4

-1.3
























Factorial axes 0 -

Figure 4.2. First factorial plane of the Correspondence Analysis of the field data (occurrence of 78 Syrphid species in 13 sampling units of the Allier (AI to A5) and Loire (L0 to L4) floodplains). The thick lines join the four main sampling units of each transect.

4.3.1.2. Reciprocal ordination of the species and their attributes

34 In the microhabitats and traits files, several data transformations are necessary before the ordination can be performed. The numerical system based on four values (0 to 3), adopted in order to describe the degree of association between the species and the categories of the variables, allows rapid transcription of available knowledge by the expert. However, in such data bases the relevant feature is provided by the relative distribution of the information among the categories of each variable. For example, for the species Eupeodes latifasciatus and the four categories of the variable "duration of the development phase", the sequence of values in the traits file was: 1 / 2 / 1 / 0. For purposes of ordination this sequence was transformed to the equivalent form: 0.25 / 0.50 / 0.25 / 0. This transformation sets the variations of the fuzzy codes between 0 and 1 and makes them add up to 1 per variable for each species. In the case of missing information for one species and one variable, the average profile of all the species for this variable can be ascribed to the species not documented. That species then plays no role in the calculation of this variable’s weight in the ordination procedure (Chevenet et al., 1994). This treatment should be limited to a very restricted number of instances .

Following these transformations, the microhabitat and trait file data are subjected to the "Fuzzy Correspondence Analysis" described by Chevenet et al., (1994), which can be regarded as a Correspondence Analysis of the data expressed as percentages per variable, or as an extension of Multiple Correspondence Analysis for use with fuzzy data. As in the case of these classical ordination methods, the analysis provides reciprocally i) species scores, which maximize the discrimination of the categories of the variables, and ii) category scores, which maximize the discrimination of the species.

Fig. 4.3a provides a picture of the ordination of the species along the first two ordination axes of the Fuzzy Correspondence Analysis. The decrease of the eigen values indicates that the first four ordination axes describe the most significant part of the total information, according to the principle proposed by Diday et al. (1982). Along the first ordination axis, the categories of the variables "microhabitat", "inundation tolerance" and "food type" are well separated, each variable explaining more than 80% of the total variance. The variables "migratory status", "microhabitat" and "food type" also contribute to a large extend to the second and third axis ordinations with more than 50% of the explained variance. The categories of the variables "number of reproduction cycles" and "overwintering phase" are the least well separated by the four axes.

35 Larval microhabitat Inundation tolerance F2

9 4 31 11 10 F1 4 1 2 3 6 7 2

8 5

Food type of larvae Number of reproduction cycles per year

4

3 1 3 2 1

2

Migratory status Overwintering phase

3

3 2

1

1 2

Eigen values: 0.51-

F1 2.2

-1.3 1.4





F2

-1.3

Factorial axes

0 -

Figure 4.3a. First factorial plane from the Fuzzy Correspondence Analysis of the fuzzy-coded trait data for 78 Syrphid species caught during the sampling programme on the Loire and Allier riverine wetlands. The 78 species (small squares) are grouped according to the categories of the six microhabitat and trait variables shown in Fig. 4.3b.

36 VARIABLES CATEGORIES SPECIES: 1 2 78 102 Microhabitat (larvae) 1 tree, free on foliage 1 2 tree, overmature, senescent 3 3 shrubs 2 4 on low growing plants, above ground 1 5 in low growing plants, above ground 2 6 litter / grass root zone, free 7 litter / grass root zone, in wood 1 8 litter / grass root zone, in bulbs 2 9 herbivore dung 2 10 water-saturated sediment, debris 3 2 11 aquatic, submerged sediment, debris 2 11 aquatic, submerged plants Inundation tolerance (larvae) 1 short respiratory tube, non tolerant 3 3 2 short respiratory tube, tolerant 3 medium respiratory tube 3 4 long respiratory tube 3 Food type (larvae) 1 microphagous 3 3 2 living plants 3 3 living animals 3 No. of reproduction cycles per year 1<1 1 21 1 3 3 32 3 3 4>2 Migratory status (adult) 1 not known to migrate 3 3 1 3 2 recorded migrant 2 3 strongly migratory Overwintering phase 1larva 3333 2 puparium 3 adult

Fig. 4.3b. Table of the 6 variables and their categories from the Microsite features and Traits files, used in the analysis. The categories are numbered as in Fig.4.3a.. Species: 1-Baccha elongata, 2-Brachyopa scutellaris, 78-Eristalis interrupta, 102-Cheilosia ahenea

37 Two groups of species are evidenced along the first axis: - On the right-hand side, the species' larvae inhabit either decaying wood (categories 2 and 7), water saturated matter (categories 10 and 11), or herbivore dung (category 9). They exhibit long or medium respiratory tubes as larvae and can therefore be regarded as inundation tolerant. They are also microphagous. - On the left-hand side of the first ordination axis, the species' larvae live either free on plant material (categories 1, 3, 4, 6), or within living plants (categories 5 and 8). They exhibit short respiratory tubes and feed on living plant or animal material.

The species belonging to these two groups are scattered along the second axis, which was associated with a gradient of increasing migratory capability and number of generation per year.

4.3.1.3. Simultaneous ordination of two matrices

The second step in the procedure looks for the common structures between the distribution of the species as sampled on site and the coded information about some of their microhabitats or traits. This can be achieved using the method of inter-battery analysis introduced recently by Chessel and Mercier (1993) and Doledec and Chessel, (1993b). This method, also presented as "Co-inertia analysis" (Dolédec and Chessel, 1994) allows the simultaneous ordination of two data matrices sharing the same set of lines. It calculates co-inertia axes, maximizing the co-variance of the factorial scores generated in the separate ordinations of the two input files. It provides therefore an ordination of the common structure of the two data sets, which maximizes simultaneously i) the variance of the factorial scores from the two separate tables, and ii) their correlation. The co-inertia analysis generates factorial scores which can be used for graphical displays as in standard ordination methods.

The co-inertia analysis looks for relationships between the distribution of the species records among the sampling units and elements of their microhabitats and traits. In Fig.4.4 it can be seen that the eigen values of the co-inertia ordination single out the first ordination axis as being generally predominant. A test of the significance of the common structure obtained may be used to compare the eigen values actually observed in the co- inertia analysis with the eigen values generated from 80 similar analyses derived from

38 random permutations of the lines of the matched tables. The results of this test, shown in Fig.4.5, demonstrate that in this case the eigen value observed for the first axis of the co- inertia analysis is significantly higher than those obtained in the random permutations, but that this is not so along the second axis. Only interpretation of the first axis is thus considered here. Along this axis, the correlation between the two matched tables is 0.63 and the explained inertia represents 94% of the inertia of the field data table and 72% of the inertia of the trait data table. The largest contribution to the ordination obtained along the first axis is the information described along the first axis of each of the previous separate ordinations.

The major output of this analysis is demonstration of the existence of a direct relationship between the ordination of the sampling stations and some of the microhabitats and traits of the species found at each sampling station (Fig. 4.4). The first axis (F1) of the co- inertia analysis discriminates the forested (A1, A2, AI) and river marginal (A0) sites of the Allier from their Loire counterparts (L1, L2, L0). When observed in relation to microhabitats, this separation can be seen to be associated with a major subdivision of the species preferences. On the left-hand side of the F1 axis, the Allier units are associated with species whose larvae are wood- and plant-tissue miners. These microhabitat categories are not or under-represented on the right-hand side of the F1 axis, where syrphids with free living larvae are mostly encountered in the units. This microhabitat segregation along F1 is also associated with different types of larval food (plant-feeding and microphagous larvae associated with miners and aquatic microhabitat categories, animal-feeding larvae mostly free-living). The third active variable along this F1 axis is inundation tolerance. Among the three other variables, one was totally non active here (migratory status), while two others (number of reproduction cycles and overwintering phase) contribute through their sparsely represented categories, which single out the more extreme units along the F1 axis. This is the case for species overwintering as puparia (Cheilosia impressa, Eupeodes luniger) and having less than one reproduction cycle per year (Temnostoma vespiforme, T. bombylans, Brachyopa scutellaris) in A1 and A2; for species overwintering as adults (Scaeva selenitica, S. pyrastri, Eristalis tenax) in L0 and L3. It can be seen from this example that simultaneous ordination, by means of co-inertia analysis of species distribution among sampling stations and some of their traits and microhabitats data, can make possible a functional interpretation of the differences between sampling stations, not solely based upon variations in their species composition.

39 In this process, as evidenced in Fig. 4.4, the actual species composition of the observed fauna of each station is hidden, but nonetheless serves as a link to match the two data sets.

AO F2

0.09 -

F1

Eigen

A4

values


F2

Factorial axes

L2a

0 -


A5 LO F1 L2 A3 L3

L4 AI L1 .5 A1 -.7 .4 A2 -.3

water-satur. aquatic sediment dung medium tube long tube in bulbs short tube tolerant on plants shrubs free in litter short tree foliage tube non overmature tolerant tree in wood (litter)

in plants Larval microhabitat Inundation tolerance

microphageous 2 >2

animals 1

plants

<1 Number of reproduction Food type of larvae cycles per year

not adult migratory strongly larva migratory recorded migrant

pupae Migratory status Overwintering phase

Figure 4.4. First factorial planes of the co-inertia analysis (occurrence of 78 Syrphid species in 13 sampling units vs. ordination of the same species on the basis of six biological traits).

40

number 14

of F1

12

values

10

8

Observed















































6

eigen value =

4
0.095

2

0

.03





.04 .05 .06 .07 .08 .09 number 25





eigen values

Observed of F2

eigen value = values

20 0.037

15

10

5

eigen values




































0





.03 .04 .05 .06 .07 .08 .09

Figure 4.5. Test of the significance of the co-inertia analysis. The eigen values of the first (F1) and second (F2) axes of the co-inertia analysis are compared with the distribution of eigen values generated by 80 similar analyses with random permutations of the lines of the matched tables.

This example uses data from individual sampling stations. But an improvement in the functional interpretation of the results would be expected if species representation had been considered in terms of habitats present on site, irrespective of the sampling stations, thereby avoiding any difficulties caused by the high mobility of adult syrphids in interpreting results from individual sampling stations. This approach would entail use of the Macrohabitats file, together with the list of the habitats observed on site.

41 Chapter 5. APPLICATIONS OF THE DATABASE

One prime objective in development of the database has been to provide a tool for use in the process of assessing site quality or site management requirements, in relation to nature conservation.

Only within the last 25 years have serious attempts been made to incorporate invertebrates into the nature conservation process. The significance or otherwise of the invertebrates of European sites designated for protection of fauna and flora has, in most cases, been disregarded during processes of site selection. But now that the concept of conservation of biodiversity draws attention to the fact that the major proportion of Europe’s biodiversity is its invertebrates, more attention is being focused on inventorisation of invertebrate faunas of protected areas. This at least makes it possible to consider the needs of parts of the invertebrate fauna when management plans for protected sites are being designed or revised, even if it is now frequently too late to ensure that the quality of the invertebrate fauna plays a role in prioritising sites for protection, or deciding where the boundary of a protected site should be.

5.1. Application of the FAEWE procedure for assessment of the “ecosystem maintenance” function of a site

A formal procedure for use of the database in site evaluation was established during course of the FAEWE project. This is used here as an example of site evaluation methodology employing the database. The FAEWE procedure dealt equally with two other invertebrate groups employed in the project and the results from the use of all three invertebrate databases are shown here, to indicate how results from syrphids may be integrated with and compared with results from other invertebrates. A potential advantage of employing invertebrates in environmental interpretation/evaluation work is the diversity of taxonomic groups available: careful selection of a small number of taxonomic groups providing complementary information can enable a wide variety of deductions to be made and establish a sounder basis for extrapolation, than can be obtained from use of one taxonomic group in isolation. Suggestions on the constitution of a cadre of taxonomic groups to use in this way are made by Speight (1986a).

42 A refinement of the use of the database employed in the FAEWE procedure was that it incorporated recognition of certain habitats as typical of active river floodplains (or river- marginal-wetlands: RMWs), so that consideration of the observed fauna of a site could be carried out not only within a framework provided by the fauna predicted for the habitats represented on-site, but also within a more restricted framework provided by which RMW-typical habitats were represented on-site. This refinement can be useful, since it carries with it the implication that if certain of these predicted RMW-typical habitats are absent from a site under investigation, then their associated species would be expected to be absent, too. This process of predicting which habitats should occur on a site can be extended beyond floodplain systems and is touched upon again in section 5.2.1.

Site evaluation may be carried out without recourse to any formal procedure like that used in the FAEWE project, but nonetheless inevitably follows an analogous step-wise process. Whatever ingredients are used in an evaluation process they require to be clearly stated, so that the logic employed can be understood.

There is not, and could never be, any absolute measure of site quality. Site quality can only be assessed through imposition of some human valuation system. It has recently become popular to choose from among various ecological approaches in selecting a valuation system to use for nature conservation purposes. But there is little stability in the resultant valuation systems, because fashions in ecology change and the products of using different attributes of ecosystems for evaluation cannot be easily compared. This quagmire may be circumnavigated by using the regional list as a basis for prediction of a site fauna, and comparing the list of observed species with the list of predicted species, so long as some standardised method can be used to calibrate the results of this comparison. In the FAEWE procedure, figures of 40% representation and 50% representation of the predicted species, on the observed species lists, have been used as a basis for decision- taking. In general, representation on-site of 50% of the species predicted for a particular natural/semi-natural habitat can be taken as indicating a reasonable representation of the fauna of that habitat on that site, in our experience, while figures higher than 50% can be used to identify sites of exceptional quality. There are no agreed international standards for recognising the quality of a site for nature conservation, whether based on use of invertebrates as tools, or on other criteria. Recommendations have been made, seeking to derive an agreed basis for recognition of sites as important at the international level for conservation of invertebrates (Speight et al, 1992), and have been followed here.

43 The format of the FAEWE site evaluation procedure based on macroinvertebrates is designed to facilitate its integration with other components of the FAEWE functional analysis procedure. At each stage of progress through the “decision tree”, the results of following the procedure with the invertebrate data from the French and Irish sites examined during course of the FAEWE project are given, in tabular form. The ecosystem function addressed by the procedure was ecosystem maintenance, as manifested through biodiversity.

5.1.1. The functional assessment procedure (FAP) Function: Ecosystem maintenance Process: Maintenance of Biodiversity

Definition: biodiversity Biodiversity is understood not only as the diversity of taxa, but also as the diversity of life strategies represented by species assemblages (e.g. modes of reproduction, feeding, dispersal,...). Because of their intrinsic habitat patchiness and environmental fluctuations, river marginal wetlands are potential sites for the coexistence of species with highly contrasted life strategies. Definition: macrohabitat and microsite feature The CORINE system set up by the EC can be used as a starting point for identifying invertebrate macrohabitats. However, the recognition of macrohabitats categories for macroinvertebrates cannot be based solely on phytosociological criteria (see Speight et al, 1997a). Therefore, a list of invertebrate macrohabitats categories has been established for use in the macrohabitats files of the database, based on CORINE categories, but incorporating additional categories important for the invertebrates covered. Furthermore, the extent to which each macrohabitat category is typical for River Marginal Wetlands (RMWs) has been coded. Within each macrohabitat type, processes such as local hydrological and microclimatic fluctuations, vegetation dynamics, local variations in sediment or debris accumulation and erosion lead to a potentially high availability of diverse microsites, which are called microsite features for the purpose of this study. Definition: RMW-Typical Macro-habitats A natural or semi-natural category occurring in the Macrohabitats files of the database is regarded as typical of river-marginal wetlands if it occurs on naturally-flooding river floodplains as a product of the dynamics of river-flow, particularly within braided and meandered stretches (the potamal stretches) of larger rivers. The database is not tuned to operate specifically for brook floodplains (floodplains in the crenal or rhitral stretches of rivers). Entirely man-made habitats introduced into functioning floodplains

44 (e.g. plantations of Salix alba) are not included as RMW-typical. A list of RMW typical macrohabitats valid for floodplain situations developed in the Atlantic and Continental zones of Europe is provided as Appendix 4 to this text. In other biogeographical zones additional macrohabitat categories could be recognised as typical for natural floodplains. Thus in the northern European zone categories of coniferous forest would be recognised as RMW typical. Definition: typical microsite features on-site RMW typical macrohabitats : Microsite features which occur naturally on RMW typical habitats during either unflooded or flooded conditions are regarded as typical. They are described in terms of their structural attributes and may be either authochthonous or allochthonous in origin. A list of them is given in Appendix 5 (an Excel file) to this text. Definition: RMW-Typical Species In the database, a species which is coded 2 or 3 for an RMW-Typical Macrohabitat is regarded as an RMW-Typical species.

Controlling variables (Cvs) and background rationale CVHabitat "faunal completeness" The occurrence in a habitat type of the species, or functional groups of species, that can be expected given their availability in the regional species pool, is considered as a sign of biodiversity maintenance. The difference between the fauna actually present and the expected one is a measure of the "functionality" of this habitat.

Information required 1- Regional list of species. The region has to be defined according to the scale and purpose of the investigation (e.g. catchment, administrative region, country, EU,...) 2- Macrohabitat and microsite feature preferences of the species in the regional list (macrohabitats and microsite features defined according to the categories defined in the Glossaries) 3- List of on-site Macrohabitats and their identification as being typical or not for RMWs in the assessment region. 4- List of species recorded on site 5- For faunistic groups with marked seasonal occurrence, phenology of the species in the regional list. 6- Threat status of the species in the regional list. 7- Regional list of RMW typical species

45 8- List of on-site microsite features and their identification as being typical for on-site RMW typical macrohabitats

Source of information 1- Published faunistic lists, experts, museum collections (Desk) 2- Expert knowledge, published information, existing data bases (Desk) 3- Site survey (Desk/Field), list of typical RMW Macrohabitats and associated typical microsite features (Appendices 4 and 5) 4- Already existing faunistic survey or faunistic survey to be carried out. In both cases ensure compatibility with the methodological guidelines indicated in the methods manual. (Desk/Field) 5 and 6- as 2 7- Regional list of floodplain typical species to be derived from regional species list and existing data base of species macrohabitat preferences, used in conjunction with the list of typical RMW habitats

Identification The completeness of a Macrohabitat fauna is assessed from the difference between expected and observed faunas. This assessment requires several steps: 1- Within the regional list of species, extraction of the species associated with on-site macrohabitats, according to the macrohabitat preference data base 2- For species with distinct seasonal occurrence, extraction of the species occurring at the time of the on-site faunistic survey(s). 3- Comparison of the predicted list obtained in 1 or 2, with the list of species recorded on-site. This can be achieved through calculation of the percentual representation of the list of species predicted for each on-site macrohabitat. Interpretation of these values is dependent upon the purpose of the assessment. 4- If data are available about the threat status of the species, the comparison can be supplemented by consideration of the threatened species associated with each on-site macrohabitat. 5- This procedure can be used not only to assess the present biodiversity of the on-site fauna, but also to anticipate its modification by known impacts that would modify on- site macrohabitats in predictable ways.

46 Tentative decision tree The following decision tree may be followed for each faunal group separately. Q1If you wish to consider the representation of all RMW-Typical species on the site, go to Q2 If you wish to consider the representation of only endemic and/or threatened species, go to Q5 If you wish to use both options, proceed through questions Q2-Q4 and then through questions Q5-Q8 Q2At the site scale, consider the percent representation of the regional list of RMW- typical species if > 40% of the predicted RMW-Typical species occurring in the region studied are observed on-site: high overall contribution of the site to biodiverrsity maintenance for the taxonomic group investigated. whether or no 40% of the predicted RMW-Typical species occurring in the region studied are observed on-site, go to Q3 for estimation of contribution to biodiversity maintenance of macrohabitats.

CLONLTBRDECZAPRE Mollusca48%45%38%36% Carabidae23%26%-- Syrphidae28%29%34%38%

Molluscs: On both Irish sites RMW-typical species reach a level of representation indicative of a high contribution to biodiversity maintenance, contrasting with the two French sites, neither of which reach that level. Syrphids: Low values are obtained from all 4 sites for syrphids in Q2, despite the high values obtained for nearly all on-site macrohabitats in Q3, because the maximal diversity of syrphid faunas is attained in deciduous hardwood forest, of which alluvial hardwood forest is an example, and all alluvial hardwood forest macrohabitats are missing from the sites studied. In other words, most RMW-typical syrphid species are associated with alluvial hardwood forest macrohabitats. Since these macrohabitats are absent from the sites studied the maximal faunal diversity of syrphids which could occur there is less than 50% of RMW-typical syrphid faunas.

The results from Q2 demonstrate an omission from the structure of the existing decision tree which could easily be made good. At present, the proportion of the entire, regional RMW-typical fauna occurring on a site is calculated (in Q2), but the next logical step is missing, namely calculation of the proportion occurring of the RMW-typical fauna of the

47 complete set of on-site RMW-typical macrohabitats, taken together. For syrphids, the existing Q2 essentially provides an overview of the condition of the investigated floodplain system in general, since it is almost inconceivable that a site could possess more than 40% of the entire regional RMW-typical syrphid fauna without being situated in a functioning, more or less complete floodplain system. As proposed above, the missing question would address the general condition of the sub-set of RMW-typical macrohabitats occurring on the site(s) investigated, in this way providing an overview of the site in general.

Q3At the site scale, consider the percentual representations of the predicted list of species in each on-site RMW -Typical macrohabitat - for each RMW -Typical macrohabitat >50% = high contribution by that RMW- typical macrohabitat to biodiversity maintenance - for all Macrohabitats go to Q4 for estimation of contribution to biodiversity maintenance of individual microsite features

Site LTBR CLON Syrphidae Mollusca Carabidae Syrphidae Mollusca Carabidae 15 Alluvial forest 30% 39% 35% (gen.) 151 Softwood(gen.) 43% 29% 1514 Gallery Softwood 48% 15141 Gallery Softwood overmature 56% 15142 Gallery Softwood mature 75% 15143 Gallery Softwood saplings 70% Scattered 0% trees/Salix/mature 21 Tall herb 70% 42% 46% 61% 46% 40% communities 2312 unimp. grass. 36% 40% 23121 unimp. grass. eutroph.(gen.) 70% 57% 64% 64% 231212 unimp. grass. eutroph.floode 75% 69% d 64 Reed/tall sedge 55% 77% 33% beds (gen.) 641 Reed beds 64% 82% 665 Running water edge 100% 80% 6651 River bank 26% 23% 713 Temporary pool 62%

48 DECZ APRE

Syrphidae Mollusca Syrphidae Mollusca 15 Alluvial forest (gen.) 48% 43% 49% 47% 151 Softwood(gen.) 70% 40% 68% 50% 151a Softwood(gen.) overmature 68% 65% 151b Softwood(gen.) mature 73% 71% 151c Softwood(gen.) saplings 92% 90% 1514 Gallery Softwood 15141 Gallery Softwood overmature 15142 Gallery Softwood mature 15143 Gallery Softwood saplings Scattered 78% trees/Populus/overmature Scattered trees/Salix/overmature 100% 100% Scattered trees/Salix/mature 100% 100% 21 Tall herb communities 55% 50% 60% 47% 23112 unimp. grass. dry, no stones 76% 21% 23121 unimp. grass. eutroph.(gen.) 70% 67% 60% 67% 231212 unimp. grass. eutroph.flooded 68% 66% 64 Reed/tall sedge beds (gen.) 54% 50% 641 Reed beds 56% 43% 642 Tall sedge beds 47% 70% 6611 water edge / unvegetated mud 67% 6612 water edge / vegetated mud 80% 80% 662 water edge / sand 100% 100% 664 water edge / standing 86% 86% 665 water edge / running 83% 83% River edge, sand/gravel 83% 83%

Molluscs: Despite the result produced by Q2, the two Irish sites show representation of less than 50% of the expected species for 2 of the RMW-typical macrohabitats represented on-site, namely softwood alluvial forest and tall herb communities. However, the other 4 achieve more than 50% representation, indicating a high contribution to biodiversity maintenance. The two French sites also show most on-site RMW-typical macrohabitats represented by more than 50% of the predicted species, which might not be expected given the outcome to Q2 for those sites. A lower contribution to biodiversity maintenance is indicated for the macrohabitat categories softwood forest, dry/semiarid uinimproved grassland without stones and reeds on the DECZ site and for tall herb communities on the APRE site.

49 Comparing representation of faunas for softwood forest on the two French sites shows that this macrohabitat is better developed on APRE than on DECZ. Character and structure of the unimproved, dry grassland without stones macrohabitat on that part of the DECZ site subject to flooding does not allow occurrence off all the species associated with this macrohabitat, a factor which must contribute to the poor representation of its associated fauna. Reeds on the DECZ site occupy a relatively small area, which may be the reason that not all species associated with large reed beds occur there.

It is noticeable that a low representation of the species predicted for tall herb communities was observed on both French and Irish sites. It is probable that this macrohabitat category is too heterogeneous for molluscs, as defined at present.

Syrphids: The significance of the RMW-typical syrphid fauna of alluvial hardwood forests is again indicated by the Q3 results, in that the level of faunal representation attained for alluvial forest in general, on the sites where some form of alluvial forest macrohabitat could be observed, was in each case below 50%. This said, the level of representation of the syrphid species typical for on-site RMW-typical macrohabitats was otherwise almost universally above 50%, indicating a reasonable level of function on all sites studied, in respect of biodiversity maintenance. One exception is the result for alluvial softwood gallery forest on the LTBR site. The explanation may be sought in the results for Q4, which show that, while the fauna associated with the herb layer of alluvial softwood forest is reasonably represented, that associated with the trees (live or dead) themselves is extremely poor, indicating they are now playing little direct part in maintaining biodiversity. The same is true of the scattered trees present on the CLON site. The only other on-site RMW-typical macrohabitat failing to attain a 50% level in its representation of associated syrphid species is the tall sedge-bed category on the APRE site. The Q4 results show that, while the vegetation-associated species from this macrohabitat are mostly reasonably represented, all groups associated with water and water-sodden ground in this macrohabitat are not. It would have been interesting to compare these results with their equivalents for the molluscs, but since more than 50% of the molluscan fauna of the tall sedge-bed macrohabitat was observed on the APRE site (Q3), its disposition between microsite features was not investigated in Q4.

Q4For each RMW-Typical Macrohabitat shown in Q3 to be represented on-site by <50% of the predicted species, consider the species associated with its typical microsite features: 50 - for each microsite feature for which >50% of associated, predicted species were observed: high contribution of this microsite feature to biodiversity maintenance - for each microsite feature for which <50% of predicted species were observed: consult specialist if this result requires further investigation.

The following tables list for the macrohabitats with less than 50% of predicted species observed, which typical microsite features have 50% or more of associated predicted species:

MOLLUSCS-French sites macrohabitats DECZ APRE 151.Softwood forest A11225 strand-line debris A31 wet mud/ooze A112172 under low growing plants, sparse A111312 on/in low growing plants A111313 on/in tussocks A11111 Foliage A11152 fallen, timber A112222 leaves,among/under forest litter 21.Tall herb communities A111313 on/in tussocks A11131on herb layer plants A11223 among/under herb layer litter 23121.no stones dry/semiarid A111311 on tall strong herbs unimproved grassland 641.Reeds A11223 among/under herb layer litter A22112 submerged mud/ooze A22213 below surface emergent water plants A31 wet mud/ooze A11214 among reed/tall sedge beds A33 sodden plant debris A111312 on/in low growing plants

51 SYRPHIDAE French sites macrohabitats DECZ APRE 15. Alluvial forest (gen) A111151 tall shrubs/ bushes/ A111121 trunk cavities saplings A11113 mature trees A111152 low shrubs/ bushes/ A11114 understorey trees saplings A111151 tall shrubs/ bushes/ A1112 upward climbing lianas saplings A11311 tall strong herbs A111152 low shrubs/ bushes/ A11312 low growing plants saplings A11311 tussocks A1112 upward climbing lianas A11151 standing timber A11311 tall strong herbs AA112171 dense low growing A11312 low growing plants plants A11151 standing timber A112221 woody surface debris + A111521 fallen timber with bark leaves A111522 fallen timber no bark A112222 dead leaves A11153 stumps A11223 herb layer litter AA112171 dense low growing A113 nests of social insects plants A1213 bulbs/ tuber A112221 woody surface debris + A1215 rotting tree roots leaves A31 wet mud/ ooze A112222 dead leaves A331 sodden timber A11223 herb layer litter A332 sodden twigs A113 nests of social insects A1213 bulbs/ tuber A1215 rotting tree roots A31 wet mud/ ooze A33 sodden plant debris (general) A331 sodden timber A333 sodden non-woody plant debris 642 Tall sedges A111 on/in plants (gen) A1113 herb layer (gen) A11131 on herb layer plants (gen) A111312 low-growing plants A1122 among/under surface debris A11223 herb layer litter A121 root zone (gen)

MOLLUSCS - Irish sites macrohabitats CLON LTBR 151.Softwood forest not represented A11225 strand-line debris A11234 on sand 21.Tall herb communities none _50% none _50% 23121.Eutr.humid/flooded A111312 on low growing none _50% unimproved grassland plants 641.Reeds A11214 among reed/tall sedge not represented beds 665.Running water edge A11225 strand-line debris A33 sodden plant debris A33 sodden plant debris A11225 strand-line debris 713 temporary pool not represented A33 sodden plant debris

52 CARABIDAE - Irish sites macrohabitats CLON LTBR 15.Alluvial forest not represented A11212 liana-covered bushes A11235 mud/ooze 21.Tall herb communities none >50% none >50% 2312.Humid or flooded A11215 grassland vegetation none >50% unimproved grassland 64.Reed/tall sedge beds A112172 sparse low growing not represented plants A11234 sand A11235 mud/ooze 6651.Riverbank A11212 liana-covered bushes A11212 liana-covered bushes

SYRPHIDAE Irish sites macrohabitats CLON LTBR 15. Alluvial forest (gen) A111152 low shrubs/ bushes/ saplings A1112 upward climbing lianas A11311 tall strong herbs A11312 low growing plants A11311 tussocks AA112171 dense low growing plants A112221 woody surface debris + leaves A112222 dead leaves A11223 herb layer litter A113 nests of social insects A1213 bulbs/ tuber A31 wet mud/ ooze A33 sodden plant debris (general) A333 sodden non-woody plant debris 151. Softwood alluvial forest (gen) A111121 trunk cavities A111152 low shrubs/ bushes/ saplings A1112 upward climbing lianas A11311 tall strong herbs A11312 low growing plants A11311 tussocks AA112171 dense low growing plants A112221 woody surface debris + leaves A112222 dead leaves A11223 herb layer litter A113 nests of social insects A1211 grass roots A1213 bulbs/ tuber A1214 stem bases A1215 rotting tree roots A31 wet mud/ ooze A33 sodden plant debris (general) A333 sodden non-woody plant debris

53 Molluscs: In the gallery softwood forest macrohabitat of LTBR 8 of a total of 35 typical microsite features harbor >50% of predicted species. This macrohabitat is not functioning, and its typical microsite features are hardly developed or missing. Two from a total of six typical microsite features of the tall herb communities macrohabitat have >50% of the predicted species on both Irish sites.

In the softwood forest macrohabitat of the DECZ site, eight from a total of 35 typical microsite features harbor >50% of the predicted species. Two from a total of six typical microsite features of the tall herb communities macrohabitat have >50% of their predicted species on site APRE. Both microsite features belong to the same categories as are on the Irish sites. Only one from 11 typical microsite features of the dry/semiarid, unimproved grassland without stones macrohabitat contributes acceptably to maintenance of biodiversity. This is one explanation for the general low level of functioning of that macrohabitat, indicated in Q3. The majority of typical microsite feature of the reed beds macrohabitat contribute to maintenance of biodiversity.This confirms indirectly the assumption that low representation of predicted snails is caused by the small area of reed beds observed on the DECZ site.

Syrphids: The significance of the absence of alluvial hardwood forest macrohabitats from all sites studied, in respect of the syrphid fauna, has already been referred to. The results from examining in more detail, in Q4, the under-representation of alluvial forest species on the sites studied highlights the significance of certain microsite features in maintaining the biodiversity of the alluvial forest syrphid fauna. In particular, the role of microsite features dependent upon overmature trees and timber is demonstrated. On the French sites, the syrphid fauna of young and understorey trees, shrubs and non-woody plants is at an observably satisfactory level of representation, but the fauna of overmature tree/timber microsites is in general not, and contributes in large measure to the low levels of representation of alluvial forest syrphids on these sites, as indicated in Q3. The fauna of timber and hollow trees on the APRE site is one exception to this generalisation, and the fauna of rotting tree roots provides another. On the LTBR site, it is only the low- growing shrubs and non-woody vegetation which is playing a satisfactory role in maintaining a fauna in the rudimentary alluvial softwood gallery forest present on the site.

54 The only non-alluvial forest category attaining less than 50% of its expected fauna is the tall-sedge bed category on the APRE site. Examination of the microsite feature results shows that for this macrohabitat, the vegetation components are generally contributing satisfactorily to maintenance of biodiversity, with the exception of tall strong herbs.

One somewhat misleading consequence of attempting to code species from all three taxonomic groups for the same categories is manifest from the syrphid results for Q4, which may be illustrated by reference to the microsite feature “upward-climbing lianas”. This category has no known syrphid species particular to it and the only syrphids coded for it are generalists, whose larvae will feed on aphids on various plant types in a wide range of different biotopes. The presence of the known syrphid fauna of upward-climbing lianas is thus of no particular ecological interest and would be expected almost wherever this microsite feature occurs - 100% representation of the associated species is not an exceptional situation. Were this microsite feature category not required for one of the other taxonomic groups involved in the project, it would not be coded for syrphids. A refinement of the database content could be achieved by careful consideration of such features to decide whether coding them adds anything which aids in interpretation. It may be wiser to leave such features entirely uncoded for taxonomic groups which have no fauna specifically associated with them, even though they may be used by generalist species.

Q5At the site scale, consider the representations of species endemic to a restricted part of Europe (localised endemic + point endemic categories in database) in each on-site typical RMW habitat for each RMW -Typical habitat presence of 1 or more restricted endemic = high contribution by that RMW-typical macrohabitat to biodiversity maintenance

Molluscs: In Ireland there is only one localised european endemic snail, Ashfordia grannularis. It is not associated with any RMW typical macrohabitat. Neither of the French sites harbor any localised, European endemic mollusc species.

Carabids: For the three groups and the four sites investigated, only one carabid species, Bembidion clarcki, belonging to the category "localised endemic" was sampled on the Irish site LTBR. It is associated with macrohabitat categories Alluvial forest, and Tall herb communities.

55 Syrphids: There are no localised syrphids endemic to Europe recorded from any of the sites studied. More than anything else, this is a reflection of the small number of such species that are associated with RMW-typical macrohabitats in the Atlantic zone of Europe. Only two of the localised endemic syrphid species in the database, Chrysogaster rondanii and Sphaerophoria potentillae, might be expected to occur in RMW-typical macrohabitats and neither of these species has been recorded from either Ireland or France. In total, there are only 6 localised European endemic syrphid species recorded from the entire Atlantic zone of Europe.

Q6At the site scale, consider the number of threatened European species present, or of threatened national species present. If there is 1 or more threatened species present: high contribution of site to maintenance of biodiversity

Molluscs: Because of the tentative nature of the list of threatened European molluscs, only the list of threatened Irish gastropods has been used for the Irish sites. 17 species are regarded as threatened in Ireland. None of them have been found on site CLON. One threatened species, Succinella oblonga, has been found on site LTBR, reinforcing the high contribution of this site to maintenance of biodiversity suggested by Q2. Since there is no list of molluscs regarded as threatened in France and the existing list of gastropods threatened at European level is of questionable reliability, it has not been possible to carry out a Q6 calculation for the French sites. It follows that there are no data available to use to carry out Q7 or Q8 calculations for the French sites.

Carabids: There is no threat status information available for carabids at European level. No Carabids which can be regarded as threatened in Ireland were found on the Irish sites.

Syrphids: More threatened species were observed for the Syrphidae than for the two other groups. The following table gives the numbers of species threatened at various levels, sampled on the studied sites.

Europe: Atlant,zone France Central.France Ireland threatened threatened threatened threatened threatened SITE LTBR not relevant not relevant 1 CLON not relevant not relevant APRE not relevant DECZ 2 2 2 2 not relevant

56 One syrphid species which may be regarded as threatened in Ireland, Helophilus trivittatus, was found on the LTBR site, categorising that site as making a high contribution to biodiversity maintenance. But H.trivittatus is by no means threatened in Europe in general, so the LTBR site can only be viewed as making a high contribution to biodiversity maintenance within an Irish context, on this basis. By contrast, the DECZ site in France can be regarded as making a high contribution to biodiversity maintenance in a broader European context, through the presence there of the two internationally threatened syrphids Eristalis picea and Sphiximorpha subsessilis. By the same token, the site makes a high contribution to biodiversity maintenance in both central France and France in general.

Q7For each threatened species observed, consult database to identify Macrohabitat preferences. If there is a preferred Macrohabitat on-site: high contribution of that Macrohabitat to biodiversity maintenance

Molluscs: Succinella oblonga, observed on LTBR, is associated with the following macrohabitats: 23121 - eutroph. humid/flooded grassland, 322 - grey dunes, 324 - dune slacks, 665 - running water edge. Only the macrohabitats 23121 and 655 occur on site.

The macrohabitat associations of Succinella oblonga suggest that the eutrophicated flooded unimproved grassland and running water edge macrohabitats on the LTBR site contribute highly to biodiversity maintenance, confirming the result obtained for these macrohabitats in Q3.

Syrphids: The following tables detail the macrohabitat preferences of the three Syrphid species sampled. Helophilus trivittatus, site LTBR 23121 eutrophic humid or flooded 1 unimproved grassland brook edge in open ground 2 permanent pool in open ground 2 edge of permanent pool in open 2 ground Eristalis picea, site DECZ temporary pool under canopy 2 Spring in forest 2 Flush in forest 2 15 Alluvial forest (gen) 1 151 Softwood alluvial forest (gen.) 1

57 151a overmature 1 151b mature 1 151c saplings 1 Sphiximorpha subsessilis, site DECZ 15 Alluvial forest (gen) 3 151 Softwood alluvial forest (gen.) 3 151a overmature 3 151b mature 1 Scattered trees / Populus overmature 2

Q8For each threatened species observed, consult database to identify Microsite features (if known) associated with the preferred on-site Macrohabitat(s) : If there is a preferred Microsite feature on-site: high contribution of that Microsite feature to biodiversity maintenance.

Molluscs: The microsite features preferred by Succinella oblonga are: A11213 among tall strong herbs A11215 among grassland vegetation A11217 among low growing plants A11223 among/under herb layer litter A11236 on soil All were observed on site LTBR.

Syrphids: The following tables detail the microsite feature preferences of the 3 Syrphid species: Helophilus trivittatus, site LTBR M22 small water movement 2 A22 In surface water 3 A221 Submerged sediment/debris 3 A2211 Fine sediment 3 A22112 mud/ooze 2 A22113 organic detritus 2 A2212 Coarse sediment 2 A22125 non-woody plant debris 2 Eristalis picea, site DECZ M2 In standing water (gen.) 2 M22 small movement 2 A21 In ground water 3 A22 In surface water 2 A221 Submerged sediment/debris (gen.) 2 A2211 Fine sediment (gen) 2 A22112 mud/ooze 2 A22113 organic detritus 1 58 A22125 non-woody plant debris 2 A3 Water-saturated ground (gen.) 2 A332 twigs 2 A333 non-woody 2 Sphiximorpha subsessilis, site DECZ A111 On/in plants 3 A1111 Trees (gen) 3 A11112 Overmature/senescent trees 3 A111122 rot-holes 3 A111123 sap runs/lesions 3

The association of the threatened syrphid species Sphiximorpha subsessilis with the macrohabitat categories of overmature and mature alluvial, softwood forest indicates a high contribution of these categories to biodiversity maintenance on the DECZ site, reinforcing the result obtained in Q3 in respect of this site. The fact that in Q4 the syrphid fauna of overmature tree microsite features on the DECZ site was shown to be under- represented demonstrates also that, even when a feature may be generally “under- performing” it may still support individual species of particular significance. The presence of Eristalis picea on the DECZ site similarly re-enforces the significance of the alluvial softwood forest macrohabitat there, but focuses on the water-sodden ground microsite features as playing a particular role. Helophilus trivittatus on the LTBR site reinforces the value of the aquatic microsite features of the unimproved, humid grassland macrohabitat, whose significance was already demonstrated in Q3.

5.1.2. Salient features of the Functional Assessment Procedure.

The results presented here show how a standardised evaluation procedure may be operated and that it produces an understandable and acceptable product. The sites studied during course of the project are arguably situated along some of the most important remaining floodplains in Atlantic parts of Europe - if the fauna of such sites failed demonstrably to reach the levels of completeness necessary to indicate that the sites contributed significantly to maintenance of biodiversity, it could reasonably be concluded that it was the procedure itself that was not functioning adequately, rather than the floodplain. But a corollary to this argument is that there is equivalent need to demonstrate the procedure operates as well with dysfunctional sites, demonstrating there a failure to contribute significantly to biodiversity maintenance Examples of use of the procedure on a range of sites of different apparent quality, both on and off floodplains, are not

59 available. The example provided is simply to demonstrate one potential use of the database.

In its present form, the FAP employed here represents a stepwise investigation of the degree of function of progressively smaller entities, the first level essentially providing information on the floodplain in general, the second on the site, the third on macrohabitats on that site, the fourth on components of particular macrohabitats on the site. It is debatable whether the user requires to pursue the interrogation of the database further than the first level, in the event that a positive outcome is obtained there. But, as can be seen from the worked examples, continuing through the process provides insights into the degree of functionality of components, even on sites shown at the first level to be generally in functional condition.

It is pertinent to question which parts of an ecosystem are addressed by the taxonomic groups employed as tools here, and whether use of any one of them alone can give a sufficiently holistic picture of a site for it to be safe to extrapolate from the results obtained to the condition of the site fauna in general. Together, the three taxonomic groups cover in the order of a 2% sample of the invertebrate fauna of the terrestrial parts of a floodplain site. While clearly better than no sample of invertebrates at all, this is not a very substantial proportion of the invertebrate fauna. Further, the three groups used here were chosen, in part at least, for their capacity to provide complementary, but different, information on the condition of a site. That they provide somewhat different information is evident from the few microsite features, in particular, for which there are results obtained in common in the worked examples. Differences between the groups, in the observed degree of completeness of their site faunas, just appear as apparently contradictory results, unless they can be explained. A good example is provided by the results from Q2, in which the molluscan calculation shows the 2 Irish sites to be highly functional, but the other two groups do not. Is this because the molluscs are reflecting conditions in components of sites to which the other taxonomic groups do not respond? Is it because molluscs reflect only certain aspects of a site’s potential, whereas the other two groups provide a more holistic reflection, indicating that components of the site not tapped by the molluscs are in poor condition? Until clear answers to questions of this type can be given and the potential of the different taxonomic groups can be more clearly stated, the non-expert user of the procedure could be forgiven for being confused by results such as have been obtained here. An approach which would avoid creation of

60 confusion in this way would be to run all available taxonomic groups of invertebrates through the FAP together, as one data set. This has not been attempted with the three taxonomic groups employed here and would entail modification of the existing database structure before it could be done. Nonetheless, it remains a possibility.

In the notes on the output obtained from each question, for the different taxonomic groups, frequent reference is made to the levels used for deciding whether the degree of faunal representation indicates a high degree of function in maintenance of biodiversity. The 40% level is used for Q2, the 50% level is used for questions Q3 and Q4. Calibration of the procedure in this way is necessary for it to work, but it requires emphasising that the percentages referred to are percentages of the fauna predicted for the times of the year (e.g. end May/beginning June + first half August for the French sites) that the sites were sampled, not of the entire predicted fauna. There can be no absolute measure of biodiversity and any evaluation system introduced is ultimately subjective, however surrounded by criteria. The levels used in the FAP here are based on the best professional judgement of invertebrate specialists who have been working in the field of site evaluation, but clearly these figures are not immutable. If experience in the use of a procedure showed that the levels used were unrealistic, they could be adjusted, so long as it were clearly stated what levels were employed and the same levels were used throughout any comparison of a set of sites.

In progressing through the decision tree, there is a noticeable tendency to use the output obtained from later questions to aid in explanation of the output of previous questions. This process stops with Q4, no finer dissection of the data being possible beyond this point using the existing structure. However, the data on species traits remains largely untapped in the decision tree as presented, and there is considerable potential for gaining clearer understanding of the results by harnessing the traits data in such a way as to provide a fifth level of precision. This is probably an unending process, in that all that is really being said is that with more data more could be achieved. But with incorporation of the traits data into the decision tree the available data files would be by and large used up. If the type of approach to employment of faunas in environmental assay enshrined in the FAP used here is to develop, and become usable for answering more than rather basic questions, there is considerable need for resources to be allocated for autecological work on the target organisms - without better knowledge of them their use as tools cannot progress beyond a certain level, which is approaching all-too-rapidly. That said, the

61 existing database is proving a remarkably powerful tool and interest in its use should help to stimulate the generation of data which will increase its capacity and versatility.

5.2. Use of the database in general site management

For a site manager, a site list of birds, mammals or flowering plants has rather different implications from a site list for some group of invertebrates. For one thing, the species are usually big enough to find, unaided. For another, getting to know most of them sufficiently well to be able to recognise them is usually a practical proposition, because there are not too many of them. Thirdly, there is usually someone within reach to provide basic information about the species. Failing that, there is normally reasonably accessible relevant literature. Certainly, invertebrate species lists are not generally so amenable to interpretation. Various attempts have been made to make invertebrate lists more usable, primarily by developing systems for indexing species, so that each has a particular value for a given type of analysis. A range of examples, mostly based on Coleoptera, can be found in Eyre (1996). Decleer and Verlinden (1992) provide an example based on Syrphidae. The Syrph the Net database represents an attempt to provide a role for all recorded syrphid species in site assessment/ management procedures, rather than confining attention to some particular subgroup(s) of species, as in more traditional “bio- indicator” work. It is hoped that, armed with relevant species lists of Syrphidae, a site manager knowing little or nothing about syrphids will be able to interrogate the syrphid database as illustrated in the following pages, thus transforming syrphid species lists from collections of meaningless names into valuable information.

The site used in this demonstration is beside the Old Kenmare Road, in the Killarney National Park, Co.Kerry (Ireland) and is referred to in this text as the OKR site. Attempts are being made by the National Park Authority to improve the quality of the OKR site. It is on a north-east facing slope, at 300- 400 metres altitude. It is covered in moorland and degraded blanket bog, with areas of poorly-drained Molinia grassland maintained by deer and sheep. In the valley bottom it incorporates an oligotrophic stream flanked by a small floodplain, covered by Myrica/Schoenus bog. This stream has occasional, isolated scrub Salix along its length, augmented by a few old, planted Larix and one or two trees of Quercus, Ilex and Betula, where it passes through a steep-sided gulley. Malaise traps were installed on this site for 3 periods of 20 days during 1995: 1-21 June, 1-21 July and

62 1-21 August. Traps were installed in pairs at each trapping station, a total of 9 trapping stations being used in all.

In this instance, use of the database involves comparison between observed and predicted fauna at many levels and using many different combinations of categories, combined from different data files. The utility of any particular comparison depends upon the questions being asked of the database. In the present context two general questions will be addressed. Firstly, in what ways might the character of the site be most usefully modified in order to improve the representation of species associated with the habitats present on the site now? Secondly, in what ways might the composition of the fauna of the site be expected to change in response to introduction of other habitats there? In addressing these questions, a step-wise process is adopted, in which attributes of a site fauna recognised as demanding further investigation at one stage are further investigated at the next stage, the stages of the process being characterised by a progressive filtering of the fauna into ever-more closely defined sub-groups. In these features, this process closely resembles the process followed in the site evaluation procedure detailed in previous pages. Site components are defined in habitat and microhabitat terms, and requirements of missing species predicted for the site are identified through their biological traits.

Examination of the site fauna commences using the entire observed fauna (i.e. site species list, which is in this case based on inventory rather than a sampling programme) and the entire predicted fauna.

The first filter used with the predicted fauna is geographic, so that prediction operates within the preferred geographic context, be it international, national or regional. The first filter used with the observed fauna relates to the habitats observed on-site. Inevitably, since the adults of Syrphidae are flighted organisms, some of the observed species will be derived from habitats not present on-site, but from somewhere in the site’s vicinity. Segregating the observed species into those associated with habitats observed on-site and those not associated with habitats on-site is thus the first action performed on the site list within the computer. The observed species associated with the habitats observed may then be compared with the predicted species associated with those habitats, to establish whether there is a reasonable representation on-site of the species expected for those habitats.

63 The concept of a “reasonable” representation of the expected species is centred on the 50% level. In other words, if 50% or more of the regionally-occurring species associated with a particular habitat are present on the site, the fauna of that habitat is taken to be reasonably represented. If less than 50% of the predicted species are present, the fauna of that habitat is regarded as under-represented. Contrasting situations illustrated by 3 different habitats on the OKR site are shown in Fig.5.1. The syrphid fauna of the poorly- drained, unimproved pasture present on the site is represented by 40% of the Atlantic- zone species for that habitat, 42% of the Irish species for that habitat, 44% of the Co.Kerry species for that habitat and 60% of the Killarney National Park species for that habitat. So, only in the context of representation of the National Park fauna would the fauna of poorly-drained unimproved pasture be regarded as reasonable on the OKR site. All of the species associated with the general surface of blanket bog in the Atlantic zone are, however, present on the OKR site, so for this habitat category the site fauna would be regarded as reasonable at all four regional levels.

Bog surface 100%

90%

80%

70% Pasture Stream 60%

50%

40%

30%

20%

10%

0%

Fig. 5.1.: The OKR species associated with three different habitats, shown as a percentage of the species associated with those same habitats in the species lists for various geographical areas. Thus the first histogram column shows that 40% of the AZ species associated with poorly-drained, unimproved pasture occur on the OKR site, and the fourth column shows that these species represent 60% of the KNP species associated with this habitat. Abbreviations: AZ = Atlantic zone of Europe; IRL = Ireland; Kerry = Co.Kerry; KNP = Killarney National Park; OKR = Old Kenmare Road site.

64 It is possible to identify whether the fauna of some particular microhabitat is under- represented in a particular habitat, or whether species-representation is poor for all microhabitats associated with that habitat. The OKR site representation of species associated with various microhabitats of the poorly-drained, unimproved pasture is shown in Figs.5.2a and 5.2b. Fig.5.2a shows that predicted species with larvae living on herbaceous plants in poorly-drained, unimproved pasture are well-represented on the site, whereas predicted species with larvae living in plant stems or bulbs in this habitat are entirely absent. Examination of the biological traits of these species shows that the former group have aphid-feeding larvae, whereas the larvae of the latter group feed on plant tissues. The plant feeders are much influenced by which plant species are present, while the aphid feeders are more dependent upon vegetation structure. The flora of the Molinia grassland on the OKR site is species poor, so an absence of plant-feeding syrphids there is not surprising. Fig.5.2b shows that the representation of species with larvae inhabiting litter-layer microhabitats is more as predicted for species whose larvae live free in the litter than for species with larvae living in sub-aqueous microhabitats of decomposing vegetable debris and cow-dung. More or less the same suite of absentee species is involved in respect of both decomposing vegetable matter and cow-dung, since their larvae may use either microhabitat. Examination of their biological traits shows all are microphages/ saprophages as larvae. Since there have been no cows on this site, the grazing animals there being deer (Cervus elephas) and sheep, absence of syrphids with cow-dung- feeding larvae would be expected. However, the absence of these same species also indicates that supplies of wet, decomposing vegetable matter in general are inadequate or in an inappropriate condition for them, on the OKR site.

The same, step-wise process may be used to identify which site components are functioning most effectively for the fauna of the habitats represented on a site. The implication then being that, for these site components, existing management practices should be continued. The entire procedure can be re-run considering only threatened species, or only threatened European endemic species, or whichever target group of the fauna requires to receive separate attention.

65 On herbaceous plants 100%

90%

80%

70%

60%

50%

40%

30%

20% In herbaceous plants(stem s) In herbaceous plants(bulbs)

10%

0%

Fig. 5.2a: The OKR species associated with three larval microhabitats of poorly-drained, unimproved pasture, expressed as a percentage of the species associated with these same microhabitats in various geographical areas. Thus the first histogram column shows that more than 80% of the Atlantic zone species occurring as larvae on herb layer plants in poorly-drained, unimproved pasture, occur on the OKR site. The eight vacant columns show that none of the species with larvae living in plant tissues or bulbs, in this habitat, occur on the site. Abbreviations: as in Fig.5.1.

80% Plant litter

70%

60%

50% Wet/subm erged plant litter

Cow dung 40%

30%

20%

10%

0%

Fig. 5.2b: As Fig.5.2a, but for different microhabitats. Abbreviations as in Fig.5.1.

66 5.2.1. Use of the database in site restoration

Two potential components of site restoration activity will be considered here: replacement of missing components of existing ecosystems and replacement of one ecosystem by another.

5.2.1.1 Replacement of missing ecosystem components

Replacement of missing components of a system implies the existence of some mechanism for identifying which components are missing. Comparison between the list of species observed and the list predicted for a site can aid in identifying missing components, just as it can in identifying poorly-functioning components.Use of the syrphid database follows the same general procedure in both instances, but in identifying missing components the predicted fauna is drawn not only from the species associated with habitats observed on the site, but from all habitats typical for the ecosystems observed on the site. Thus, in the case of the OKR site, in identifying missing ecosystem components the species predicted for blanket bog would include not only those for the habitats of blanket-bog surface, flushes, springs and streams, all of

Blanket bog habitats 100% All b.bog habs.

90%

Flush 80% Stream Spring Pool edge

70%

60%

50%

40%

30%

20%

10%

0%

Fig. 5.3: The predicted Irish syrphid fauna of habitats that are typical constituents of blanket bog, compared with the observed fauna for these habitats in selected geographic areas. Thus the first histogram column shows that 70% of the stream-associated blanket-bog species known in Ireland occur on

67 the OKR site. The second column shows that the same proportion of these species occurs in the Killarney National Park in general. Abbreviations: All b.bog habs. = all blanket-bog habitats; others as in Fig.5.1. which were observed on-site, but also for pools, which were not observed. The result of comparing observed fauna with predicted fauna are shown in Fig.5.3. It is apparent from Fig.5.3 that there is, indeed, a group of syrphid species associated with pools in blanket bog in Ireland, only 45% of which are found on the OKR site, making this the poorest- represented group of blanket bog species occurring there. These histograms also show that whereas more than 90% of the Irish species in this group occur in Kerry, only just over half of them are present in the Killarney National Park. Further investigation of these missing species shows they all have larvae which are either predatory (on aphids) or aquatic/subaquatic saprophages.

5.2.1.2 Replacement of one ecosystem by another

Replacement of an ecosystem existing on a site by another ecosystem implies some method for deciding which alternative ecosystem would be the preferred option. On the OKR site, active consideration is being given to replacement of the poorly-drained, unimproved Molinia grassland by some form of forest. This proposition requires consideration of what is the potential for establishment on-site of a representative fauna of the alternative forest types. An approach to evaluating the relative merits of establishing Quercus forest or Alnus/Salix forest there is illustrated in Fig.5.4. In conducting this comparison, all species observed on the OKR site are included - the species recorded on the OKR site, but not associated with existing habitats there, may be derived from Quercus or Alnus/Salix forest somewhere in the vicinity, and would so indicate a capacity for species from these forest types to reach the OKR site. Fig.5.4 shows that the existing site fauna contains few species associated with either Quercus or Alnus/Salix forest, and that, except for species associated with scrub, the existing fauna would contribute somewhat more to Alnus/Salix forest than to Quercus forest. The fauna of the Killarney National Park in general would also have a greater capacity to contribute to the fauna of Alnus/Salix forest, were it established on the OKR site, than to the fauna of any Quercus forest established there, again with the exception of scrub fauna. The same is true of the fauna of Kerry, the County within which the Park is located. Overall then, Alnus/Salix forest established on the OKR site might be expected to have a more complete fauna than Quercus forest established there.

68

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Clearings MatureOvermature

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Fig. 5.4: Syrphid fauna predicted for selected habitats typical of two forest types, Quercus and Salix/Alnus, if established on the OKR site, based on the known fauna of these forest types in areas of different geographic extent. Thus, the first histogram column shows that c.20% of the Irish species associated with scrub Salix would be predicted to occur on the OKR site if Salix forest were established there, basing prediction on the existing OKR site fauna. The third column similarly shows that c.45 % of the Irish scrub Salix fauna would be expected, basing prediction on the fauna of the National Park in general. Abbreviations: as in Fig.5.1.

Another line of enquiry concerns the extent to which an introduced ecosystem type might be expected to support particular groups of target species, for instance threatened species. In considering this question it is necessary to address such issues as what threatened species are available for colonisation of the site, what microhabitats they occupy and for how long it would be necessary to monitor the site in order to evaluate the success of the undertaking. As an example, Quercus-forest syrphids threatened in Ireland are considered in Fig.5.5. The first set of three histogram columns shows that no species in this category occur on the site at present, that only 10% of them occur in the Killarney National Park and only 30% of them have been recorded from Co.Kerry in general. The majority of the threatened Irish Quercus forest syrphids cannot, therefore, realistically be regarded as available for colonisation of the OKR site. The second set of three histogram columns shows that most of the threatened species in this category are saproxylics, so that any Quercus forest established on the OKR site would have to be of adequate size to provide for all age classes of trees up to and including overmature and senescent trees, in order to maintain these species on-site, were they to establish themselves there. The third set of

69 three columns shows that most of these threatened species would not be expected to establish themselves on-site in less than 50-100 years, so that long-term monitoring would be necessary to find out whether establishment of Quercus forest on the OKR site led to establishment of these species. The spreadsheet used in production of this set of histograms has as yet only been coded for the Irish species and does not form part of the published version of the database.

Oak forest syrphids threatened in Ire land 80% establishm ent time

70%

microhabitat 60%

50%

40% IRL spp. in diff.lists

30%

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Fig. 5.5: Threatened Irish Quercus forest syrphid species categorised in three different ways: as represented on selected lists (three columns at left); as associated with three different larval microhabitats (three columns at centre); according to potential rate of establishment on-site (four columns at right). For explanation, see text. Abbreviations: as in Fig.5.1.

5.3. Comparisons between regional lists.

To illustrate this potential application of the database, syrphid species lists for various parts of the Atlantic zone of Europe are compared. The lists are derived from Ireland, Great Britain, North France, Central France and the Atlantic Zone in general, these parts of Europe being defined as in Speight (1996b), where this comparison was first presented.

The syrphid list used for North France comprised 228 species, while the list for Great Britain included 251 and the list for Ireland 171. The list compiled for the Atlantic zone in general comprised 345 species when this comparison was carried out. The species may

70 first be categorised according to which general habitat categories they utilise. The distribution of the species between general habitat categories is indicated in the histograms presented in Fig.5.6, with the species complement for each of the five lists being considered separately. It should be noted that because the habitat associations of species are not necessarily exclusive, some species occur in more than one of the habitat categories employed. In consequence, summing the percentages shown by a set of histograms for a particular species list does not give a total of 100%.

60%

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Series3

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Deciduous forestsConiferous forestsWetland O pen ground

Fig. 5.6: Representation of syrphids associated with major habitat categories in various parts of the Atlantic zone of Europe, expressed as the percentage of each regional list associated with each category. Example: column 1 shows that approximately 54% of the syrphid fauna of Central France is associated with deciduous forest habitats. Regional lists used: AZ = Atlantic zone, general; CFR = Central France; GB = Great Britain; IRL = Ireland; NFR = North France.

Fig.5.6 shows that the major component of the Central France fauna is deciduous forest species, and the similar preponderance of deciduous forest species in the lists for North France and Great Britain suggests this is not unusual for parts of Atlantic Europe. In the past, the natural vegetation types predominating in these parts of Europe were all deciduous forest of one sort or another (Ozenda, et al, 1979), so the dominance of woodland syrphids in these faunas is not surprising. But, given the considerable influence of man’s activities in Europe over the past 5000 years, particularly in removing forest from most of the continent, the dominance of woodland syrphids in these lists might quite reasonably be regarded as a “fossil” feature, reflecting the past rather than the present. 71 The case of Ireland, where forest removal by man has been most extreme and wetland syrphids predominate, might simply be interpreted as being as much a consequence of the land-use history of the island as of climatic influences. The increasing diversity of indigenous, deciduous forest types represented locally, as you travel from Ireland through to Central France, will also increase the capacity of Central France to support deciduous forest syrphids, in comparison with Ireland. For instance, beech (Fagus) forest is not indigenous to Ireland, whereas it is indigenous to the parts of Europe covered by the other lists used in Fig.5.6, and 5% of the deciduous forest syrphids listed for Great Britain, North France and Central France are species which occur almost exclusively in beech forest.

Fig.5.6 shows there is a progressive increase in the proportion of deciduous forest syrphids in each list as you pass towards Central France, but does not address the question of the number of deciduous forest species in these lists. Contrasting the Irish and Central France lists, Fig.5.6 indicates that there is either a lesser diversity of wetland species represented in the Central France list, or a considerable increase in the deciduous forest species in that list, as compared with Ireland. Examination of the lists themselves shows that both processes are occurring.

The general habitat grouping with the smallest component of species in the Central France list is conifer forests, in stark contrast to the situation for deciduous forests. Once again, comparison with the other species lists shows this is not an exceptional condition. Indeed, comparing the situation with the species lists for the entire Atlantic zone shows that syrphids which utilise conifer forest habitats are but a minor constituent of the entire regional fauna. Presented as in Fig.5.6, the conifer forest contingent of syrphid species includes species shared with deciduous forest, i.e. species which utilise either conifer or deciduous forests as habitat. If syrphids which use only conifer forest habitats are considered separately, an even more extreme situation is revealed. More than half of the conifer forest syrphid fauna consists of species shared with deciduous forest - the species occurring only in conifer forest comprise only 8% of the lists for Central France and North France, 9% of the list for Great Britain and 6% of the Irish list. The most extreme case is again that of Ireland. Indigenous conifer forests (of Pinus sylvestris) are believed to have been almost entirely eradicated from Ireland for 500 years or more, so the 6% of the fauna made up of strictly conifer forest syrphids is believed to be of very recent origin, namely syrphids which colonised the island subsequent to introduction of

72 commercial plantations of introduced conifers, which began on a large scale only during the present century.

The small size of the conifer forest syrphid fauna in these various parts of the Atlantic zone of Europe shows that the current practice of replacing native deciduous forests with conifers, whether the conifers are of European or other origin, must result in a net decrease in the diversity of the syrphid fauna. Furthermore, Fig.5.6 shows that these parts of the Atlantic zone are also typical of the Atlantic zone in general, in the relative insignificance of the conifer forest fauna. At a time when European nations are attempting to grapple with the problems of maintaining biological diversity, following on from the provisions of the Biodiversity Convention, the potential influence of coniferisation on the diversity of forest faunas perhaps requires to be given greater prominence. If other taxonomic groups are similar to the syrphids, forest faunas will progressively change from being the largest group of species in a local fauna to its smallest group of species, as an area’s forests are converted from deciduous trees to conifers, to judge from the information provided by the comparison of species lists presented here. Admittedly, with enormous hectarages of conifers now established outside their natural range, from the Atlantic to the Urals, some influx of conifer forest fauna from outside Europe, from Siberia, might be expected. To-date, this invasion has largely resulted in the appearance of a number of conifer pest species and their associated predators - some of them syrphids - which has in no way counteracted the impoverishment of forest faunas caused by coniferisation itself.

Fig.5.7 demonstrates that syrphids with predatory (aphid-feeding) larvae are predominant in the conifer forest fauna of the Atlantic zone of Europe, and that this dominance is expressed in its most extreme form in the Irish list. Managed, commercial conifer plantations lack both the associated ground flora and the overmature trees of indigenous, natural forest. In consequence, the syrphids associated with both ground flora and old trees are almost entirely absent in Irish conifer plantations and hence absent from the Irish syrphid list. In contrast, the presence of more extensive areas of less intensively managed indigenous forest, both conifer and deciduous, in both North and Central France, is reflected in the reduced dominance of forest syrphids with predatory larvae

73

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d.predatorsc.predatorsd.herbivores c.herbivores d.saproxylics c.saproxylics

Fig. 5.7: Representation of syrphids with predatory, plant-feeding or saproxylic larvae in the deciduous forest and coniferous forest faunas of various parts of the Atlantic zone of Europe, expressed as a percentage of the relevant forest fauna of each species list Coding of columns as in Fig.5.6; prefix c = coniferous forest; prefix d = deciduous forest e.g. dpredators = deciduous forest syrphids with predatory larvae. Example: column 9 shows that 90% of the Irish conifer forest syrphids are species with predatory larvae.

there, the species with saproxylic (dependent upon old/senescent trees) or plant-feeding (leaf, stem or root-mining) larvae becoming increasingly evident components of the fauna. The fauna for Central France again represents the opposite extreme to the Irish fauna, in this regard. The percentage of predatory species in the Central France list most closely resembles that for the Atlantic zone in general, in respect of both deciduous and conifer forests. It is noticeable that in both conifer and deciduous forest,

Syrphids with predatory larvae are the dominant component in all lists. However, even in the list for the Atlantic zone in general, the predatory species comprise a greater proportion of the conifer forest fauna than the deciduous forest fauna - in deciduous forest, the saproxylics and plant-feeding species combined represent as great a proportion of the fauna as the predators, while in conifer forest there are twice as many predatory species as the combined total for plant-feeders and saproxylics.

74

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peatlands other wetlands dry, unimproved pasturehumid, unimproved pasture

Fig. 5.8: Proportion of the Atlantic zone syrphid fauna of particular habitat types represented in various species lists, expressed as the percentage of the Atlantic zone fauna of each habitat type represented on each list. Example: column 3 shows that 90% of the Atlantic zone syrphid species associated with peatlands are represented in Great Britain. Coding of columns as in Fig.5.6.

Turning to the wetland and open ground groups of habitats, the most evident difference between the species lists lies in the high proportion of wetland species in the list for Ireland. But this is more because of an absence from Ireland of forest species represented on the other lists, than because more wetland species are present in Ireland than in the other lists. This can be recognised from considering the proportion of Atlantic zone wetland species on each list. Subdividing wetland habitats into two groups, peatland (fen and bog) and non-peatland habitats, produces the results shown in Fig.5.8, which indicates that for both sub-groups of wetland habitats a smaller proportion of the associated Atlantic zone fauna occurs in Ireland than in Great Britain, despite the fact that deciduous forest species are the dominant component of the British fauna and wetland species predominate in the Irish fauna. Another characteristic of these wetland faunas, indicated in Fig.5.8, is that peatland species are somewhat better represented in both Ireland and Great Britain than are the species associated with other wetland habitats, whereas the converse is true in both Northern and Central France. A similar relationship exists between the species complement of humid and dry unimproved pasture, which, between them, comprise the vast majority of the open ground species. Thus, species

75 associated with humid, unimproved pasture, are better represented on the Irish and British lists, while species associated with dry, unimproved pasture are better represented on the two French lists. The list for Central France represents the extreme of this condition, with the lowest representation of peatland-associated syrphids and the best representation of syrphids associated with dry, unimproved pasture. The character of the climate is presumably to a significant extent responsible for these features of the syrphid fauna.

Indubitably, a myriad influences have combined to shape the syrphid fauna of Atlantic parts of Europe as we find it today. But this overview shows that some features of regional syrphid faunas are probably in large part a consequence of the history of man’s activities in the area, just as others are most easily understood by considering parameters of the local climate. Whatever the precise reasons may be for some particular feature of a fauna, the ecological balance of regional species lists, as employed here, is demonstrably a tool of potential value in environmental interpretation.

In a second example of comparison between regional species lists, Speight (2000b) compares attributes of the regional species lists for conifer plantation syrphids from various parts of Europe, from Ireland to Switzerland. This comparison demonstrates that conifer plantations are unlikely to function as "ecological corridors" for the transport of all elements of the European syrphid fauna of Abies/Picea/Pinus forests, into/through those parts of Europe where conifer forest is not indigenous. The only element of the conifer forest syrphid fauna which seems generally able to make use of these plantations is the species which feed on conifer foliage aphids, and even these species show little capacity for rapid spread through conifer plantations. For instance, in Ireland, where none of these conifers are indigenous (Pinus sylvestris did occur in Ireland during the post- glacial, but became extinct some hundreds of years ago), only one Picea-associated species, Eriozona syrphoides, is recorded so far, although the "ecological corridor" provided by conifer plantations has now been in place for nearly 100 years.

76 Chapter 6. PROGRESS AND LIMITATIONS

A primary objective of the FAEWE project was to make progress toward establishment of an expert system, to apply in particular to functional analysis of river margin wetlands. The studies on invertebrates incorporated into the project adopted the spirit of this objective, in attempting, wherever possible, to set up and test mechanisms which would enable non-experts to gather, handle and interpret data relating to the taxonomic groups of invertebrates addressed by the project. For syrphids, selection of field techniques which would allow maximal involvement of non-experts in the sample-collection process proved reasonably easy. Ways of transferring the necessary laboratory work to non- experts proved an obdurate problem. In particular, the identification of material collected remains firmly in the hands of experts and all evidence points to this being an immutable requirement for the foreseeable future, whatever taxonomic group of invertebrates is involved. The procedure developed is entirely dependent upon identification of the target organisms to species, correctly.

At the outset of the FAEWE project there was no mechanism in existence which allowed the interpretation of terrestrial invertebrate faunas to be carried out by non-experts, other than those which treated the organisms simply as integers. The project saw the development, testing and use of a novel system which, at this point in time, can already be used by non-experts to interpret species lists for certain invertebrates, making maximal use of available biological data about the species. Indisputably, more incisive and comprehensive interpretations can be obtained by use of the system for the same purposes by specialists in the taxonomic groups concerned. However, procedures, like the FAP run through here, can be conducted by someone who has no knowledge of the organisms involved, armed just with species list and habitat data collected from the target sites.

The problems encountered during course of this work nearly all relate to the process of setting up a system whereby non-experts might interpret species lists. These problems were encountered every step of the way, right from the point in time at which it was necessary to decide which taxonomic groups of invertebrates would be the most appropriate to use as tools in this endeavour. The criteria employed in the selection process were enumerated in the Introduction to this volume. One of these criteria was

77 that adequate biological information should be available for the organisms chosen, such that it would be possible to deal with entire faunas, rather than having to restrict attention to a few so-called bio-indicators which had been studied in depth. Following decision on which groups to use it was rapidly discovered that, despite an apparent wealth of published information, the published data sources all-too-frequently lacked necessary ingredients. Much reliance then had to be placed upon consultation of experts to obtain missing data, much of which had never been published. In the case of the syrphids, this meant contacting experts in many different countries.

The files which constitute the database have undergone considerable evolution since they were first set up. The fuzzy coding system was introduced at the outset and has proved invaluable, but the categories coded were for some years in a state of almost continuous flux. Originally envisaged as paired files, one coding habitat data and the other coding biological traits data, both the number of categories of data included in the files and the number of files, increased with experimentation in their use. In large part, the proliferation of categories and files related to attempts to refine the prediction process on which depended provision of a reliable standard, with which to compare observed faunas. Initial prediction attempts resulted in over-prediction of species, so that additional data were then needed to provide filters allowing of more successful exclusion of wrongly predicted species. The outcome of this evolution was the use of the macrohabitats files as the basis for prediction of the expected fauna of a site, with the supplementary habitat categories and certain additional files, developed from the original traits file, introduced specifically to shape the prediction process. The flight period file used in predicting the syrphid fauna of a site is one such additional file. Other elements of the original traits file developed into the microsite features file, used not in prediction, but for investigation of differences between predicted and observed faunas. Similarly, it became apparent that some way had to be found to deal with threatened species as a distinct issue, since other approaches to evaluation of site faunas almost inevitably explored the use of the threat status of the observed species, even if the rest of the fauna was entirely ignored. This resulted in production of the range and status file. The relative availability of data on the different species, for coding them in these various files, has already been referred to. Suffice it to say that, even when relevant information existed, it proved difficult to encapsulate in ways that made possible the construction of a common reference system of categories for the three taxonomic groups employed on the FAEWE project. Essentially, each author or expert used their own terminology for describing habitat, habits, ecology

78 etc. Given that large projects had been in place for some years to set up Europe-wide systems for classifying habitats, it might be assumed that, for macrohabitats at least, delineation of categories to use would have been relatively simple, just by adopting the CORINE system, for instance. This was attempted, and where possible CORINE “habitat” categories are used in the macrohabitats file. But the CORINE “habitats” have little in common with invertebrate habitats (see Speight et al, 1997a) and it proved necessary to generate a complementary system of categories to “fill in” the invertebrate habitats not recognised by CORINE. In the process it was discovered that each taxonomic group required to have macrohabitat categories defined for it alone, a process which would be magnified further with addition of each supplementary taxonomic group of invertebrates one might wish to incorporate into the system. This problem proved even greater, when it came to defining microsite feature categories which were meaningful. The number of microsite feature categories not shared by the three taxonomic groups studied is greater than the number that is shared. This has produced a complication evidenced in the FAP decision tree calculations included in this text, namely that there are surprisingly few contact points between the microsite feature results for the three groups. While it might be argued that it is inevitable that organisms with such different evolutionary history and strategies would provide different types of input to such a decision tree and should thus provide different sorts of information about a site, there are circumstances in which it would nonetheless be helpful if the results from the three groups were more directly comparable!

A whole range of issues await exploration with the syrphid database, few of which have been referred to in this text. The database may be used to predict impacts upon the fauna of specified changes in macrohabitat occurring on a site/in a region, for predicting faunistic change likely to occur accompanying initiatives in restoration ecology, or for investigating many aspects of the inter-relation between groups of species and their attributes. Recently, the focus of exploration of applications of the database has switched from macrohabitats to microhabitats, with recognition of the direct link that exists between microhabitats and impact of land management operations (Speight, 2000a). This has resulted in addition of an additional file to the database, in which the degree of adverse impact of a range of animal farming management practices upon the species are predicted (Speight et al, 2000). The full range of potential applications of the database is still not defined.

79 REFERENCES

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86 Appendix 1: HABITAT SURVEY FORM

SITE NAME:

------LOCATION:

------SITE NUMBER:DATE OF SURVEY:

CODE NUMBER OF CODE NUMBERS OF ASSOCIATED HABITAT OBSERVED SUPPLEMENTARY HABITATS

NOTES

87