April 2006

ISSN 0952-7583

Vol. 19, Part 1

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BRITISH JOURNAL OF

ENTOMOLOGY

AND NATURAL HISTORY

BRITISH JOURNAL OF ENTOMOLOGY AND NATURAL HISTORY Published by the British Entomological and Natural History Society and incorporating its Proceedings and Transactions

Editor: J. S. Badmin, Coppice Place, Perry Wood, Selling, nr Faversham, Kent ME 13 9RB (Tel: 01227 752291) email: jbadmin@btinternet.com

Associate Editor: M. Wilson, Ph.D., F.R.E.S., F.L.S. Department of Biodiversity & Systematic Biology, National Museums & Galleries of Wales, Cardiff CF10 3NP. (Tel: 02920 573263) email: Mike.Wilson@nmgw.ac.uk

British Journal of Entomology and Natural History is published by the British Entomological and Natural History Society, Dinton Pastures Country Park, Davis Street, Hurst, Reading, Berkshire RG10 OTH, UK. Tel: 01189-321402. The Journal is distributed free to BENHS members.

© 2006 British Entomological and Natural History Society.

Typeset by Dobbie Typesetting Limited, Tavistock, Devon.

Printed in England by Henry Ling Ltd, Dorchester, Dorset.

BRITISH ENTOMOLOGICAL AND NATURAL HISTORY SOCIETY

Meetings of the Society are held regularly in London, at the rooms of the Royal Entomological Society, 41 Queen’s Gate, London SW7 and the well-known ANNUAL EXHIBITION is planned for Saturday 1 1 November 2006 at Imperial College, London SW7. Frequent Field Meetings are held at weekends in the summer. Visitors are welcome at all meetings. The current Programme Card can be obtained on application to the Secretary, J. Muggleton, at the address given below.

The Society maintains a library and invertebrate collections at its headquarters in Dinton Pastures, which are open to members on various advertised days each month, telephone 01189-321402 for the latest meeting news. The Society’s web site is: http://www.BENHS.org.uk

Applications for membership to the Membership Secretary: D. Young, 22 Wordsworth Close, Saxmundham, Suffolk IP17 1WF. Tel: 01728 603568.

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Society Website: www.benhs.org.uk for recent information on the Society’s meetings programme and general society details.

Cover photograph: Andrena bucephala Stephens (Hymenoptera: Apidae), a globally rare bee recorded from Warwickshire limestone quarries. Photo: S. Falk.

NOTE: The Editor invites submission of photographs for black and white reproduction on the front covers of the journal. The subject matter is open, with an emphasis on aesthetic value rather than scientific novelty. Submissions can be in the form of colour or black and white prints or colour transparencies.

Editorial Committee:

D. J. L. Agassiz, M.A., Ph.D., F.R.E.S. R. D. G. Barrington, B.Sc.

P. J. Chandler, B.Sc., F.R.E.S.

B. Goater, B.Sc., M.I.Biol.

A. J. Halstead, M.Sc., F.R.E.S.

R. D. Hawkins, M.A.

P. J. Hodge

T. G. Howarth, B.E.M., F.R.E.S. I. F. G. McLean, Ph.D., F.R.E.S M. J. Simmons, M.Sc.

P. A. Sokoloff, M.Sc., C.Biol., M.I.Biol., F.R.E.S.

R. W. J. Uffen, M.Sc., F.R.E.S. B. K. West, B.Ed.

Registered charity number: 213149

BR. J. ENT. NAT. HIST., 19: 2006

1

A SAWFLY, PRISTIPHORA LEUCOPUS (HELLEN), (HYMENOPTERA: TENTHREDINIDAE) NEW TO BRITAIN

K. J. Grearson

10 Eastfield, Ashton Keynes, Swindon, Wiltshire SN6 6PR

Abstract

Lime Sawfly, Pristiphora leucopus (Hellen) (Tenthredinidae, Nematinae), is reported from Britain for the first time. Notes on the distribution and identification of the insect are given, including a revision to Benson’s (1958) identification key.

Introduction

In July 2003 and June 2004 a number of Pristiphora larvae were beaten from limes, Tilia spp., at eight sites in Wiltshire and one in Norfolk (per John Badmin). Most of these were reared out. The host plants were Tilia x vulgaris Hayne and T. cordata Miller. The rearings produced 14 adults of a species which could not be identified using the British key (Benson, 1958). Determination was carried out by reference to Zhelochovtsev (1988). Comparison was also made with specimens of adults and larvae, collected in Finland, borrowed from the collections of the Natural History Museum, London (NHM). In 2002, 2003 and 2005 Andrew Halstead swept several Pristiphora females from limes in the arboretum at RHS Garden, Wisley, Surrey and he had not succeeded in identifying them. Comparison of these with the Wiltshire reared specimens and those from the NHM confirmed that they were all of the same species. Pristiphora leucopus (Hellen) occurs in Finland, Russia, Ukraine (Zhelochovtsev, 1988) and Germany (Andrew D. Liston pers. comm.), always in association with Tilia. Of the 49 British species of Pristiphora , 41 are believed to be monophagous (Lacourt, 1999). It is not known whether this species is a recent arrival in the country or has simply been overlooked.

Description of larva

The larva has a uniformly pale green body, with a smooth, shiny texture and a narrow whitish line along the flanks level with the spiracles. The larva is a leaf-edge feeder and the green body colour is the same colour as the lime leaves. There are six pairs of prolegs, one on each of abdominal segments 2-7, plus a pair of anal prolegs on the tenth and last abdominal segment. The young larva, as illustrated in Fig. 1 , has a very dark head and bold black coxal marks on the thoracic legs. During the final days of feeding the head pattern, as shown in Fig. 2, becomes more contrasting with a pale greenish-grey background colour and black coronal and parietal stripes. A black marking on the frons is separate from the coronal stripe. The coxal markings are less prominent in older larvae. Full-grown larvae are 15 mm long. The head and coxal markings of the reared larvae were an exact match with the NHM loan material. When the reared larvae had finished feeding they spun a brown cocoon approximately 8 mm long among the leaves, no other medium having been provided. No change was noted in the appearance of the larvae at this stage. Adults emerged a few days later. It is estimated that the cycle from egg-laying to emergence of the adults took about three weeks. The collected larvae were reared in an indoor situation and the cocoons kept indoors.

BR. J. ENT. NAT. HIST., 19: 2006

Description of adults

The reared adult females were easily assigned to the genus Pristiphora using Benson (1958). The description is as follows: body length 4.75-5 mm. The head, thorax and abdomen are predominantly black. The surface of the head and thorax is shiny, finely punctured and pilose. The surface sculpture of the tergites is coriaceous. Labrum and mandibles are brown, maxillary and labial palps yellowish-white, each of the latter having a small amount of dark pigment on the basal segments. The ocelli are positioned well back on the vertex leaving a very narrow post-ocellar area. Hind ocelli are twice as far apart as the distance between one of them and the edge of the occipital carina. There is a small pit behind each ocellus. The two basal antennal segments are black with an apical brown rim. The flagellar segments 3-9 are light brown with a narrow darker line longitudinally along the top, fading near the tip so that the apical segment is wholly light brown.

The only paler parts of the thorax are the outer two-thirds of the tegulae which are white merging into black at the interface with the mesonotum and the pronotum, which has pale hind margins adjacent to the tegulae.

The whitish cenchri (Fig. 3a) are more or less oval in shape, slightly angled so that the outer tip of each is slightly further forward than the inner tip and as far apart as approximately 1 .4 x the length of one cenchrus. The metascutellum is shiny with a few short hairs. Wings are hyaline with a covering of microtrichia, densest at the wing tips. The stigma is large, brown in the middle and surrounded by a pale border. The costa is pale brown and slightly darker at the swollen apex.

The legs are yellowish-white with a small amount of black as follows. The coxae are black basally. The pale femora have a minimal amount of black surface shading, the extent of which varies between individuals but is never the dominant colour. (Please refer to the note below regarding the femur colour of some adults which emerged in mid-May.) The apices of the tibiae and the tarsal segments are tipped with light brown. The tarsal claws all have a large inner tooth as illustrated in Fig. 3c, best described as sub-bifid. The inner hind tibial spurs are roughly half the length of the basitarsus.

Some abdominal tergites have pale medial apical markings. These are widest and palest on tergite 1 and become smaller and darker from tergite 2 onwards. Tergites 8 and 9 have no pale markings. The sawsheath, viewed from above, is illustrated in Fig. 3d. It is short, not protruding far beyond the ninth tergite and as wide at the tip as the apex of a hind tibia when viewed from above. The saw, illustrated in Fig. 3b, has rows of hairs on the back of the lines arising from each of the saw teeth from tooth 5 onwards. The hairs over tooth 5 are short, sparse and difficult to see even under high magnification (e.g. x 180).

As with the larvae, the reared adults matched the NHM loan material exactly.

The adults swept by Andrew Halstead in the arboretum at Wisley Garden during mid-May had considerably more black on the femora than any of the specimens reared from larvae of the summer generations. The hind femora of five adults collected during May at Wisley had black/white in the ratio of approximately 70:30 for an individual caught on the 10 May ranging up to about 40:60 for individuals caught on 24 May. The six dark-legged species of Pristiphora arising from couplet 7, P. bifida Hellen, P. melanocarpa (Hartig), P. ruficornis (Olivier), P. coniceps Lindqvist, P. armata (Thomson) and P. confusa Lindqvist, of Benson’s key all have a tiny amount of white on the hind femora. More than 50 examples, the majority on loan from NHM, were examined and none had more than 10% of white, most of them considerably less. Individuals of P. leucopus swept at Wisley in June and July were similar to the reared adults described above.

Fig. 1. A young larva of Pristiphora leucopus showing mainly black head and black coxal markings on thoracic legs

Fig. 2. An older larva of Pristiphora leucopus showing the head markings and reduced coxal markings of the final instar

BR. J. ENT. NAT. HIST., 19: 2006

3

Fig. 3. Pristiphora leucopus female, (a) metascutellum and cenchri, (b) saw, (c) tarsal claw, (d) dorsal view of sawsheath. (drawings by K.J.Grearson)

Fig. 4. Pristiphora leucopus male penis valve, (drawing adapted from Lindqvist, 1969)

No British male has been seen so far. Lindqvist (1969) referred to a single rearing of larvae in Finland which were overwintered successfully and from which emerged 23 females and four males in the following spring. It seems that it may be normal for males to be in the minority. Lindqvist’s drawing of the male penis valve is included here as Fig. 4.

British material examined

Ashton Keynes, Wilts. (SU048938)-6 July 2003 -Larva on Tiliax vulgaris - reared female emerged 18 July 2003.

Warminster, Wilts. (ST879442)-8 July 2003 -Two larvae on T. cor data- reared females emerged 22 July 2003?

Water Eaton, Wilts. (SU 133950) -28 July 2003 -Two larvae on T. cor data- reared females emerged 10 and 11 August 2003.

Water Eaton- 31 May 2004 -Larva on T. cor data -reared female emerged 12 June 2004.

Webb’s Wood, Wilts. (SU045855)-1 June 2004 -Five larvae on T. cor data- two reared females emerged 11 and 12 June 2004. Female swept 22 July 2005.

4

BR. J. ENT. NAT. HIST., 19: 2006

Purton Stoke, Wilts. (SU095897)-4 June 2004-Ten larvae on T. x vulgaris- Four preserved and five females emerged 12 June 2004.

Oxborough, Norfolk -4 June 2004 -Three larvae on T. cor data (J. Badmin)- Females emerged 14-16 June 2004.

Dauntsey, Wilts. (ST993819)-4 June 2004 -One larva on T. x vulgaris, released.

Purton, Wilts. (SU093875)-4 June 2004 -Two larvae on T. x vulgaris, released.

Lydiard Millicent, Wilts. (SU081860)-4 June 2004 -One larva on T.x vulgaris, released.

RHS Gardens, Wisley (TQ064577)- Females swept by Andrew Halstead on 19 June 2002, 14 May 2003, 24 May 2003 (3), 10 May 2005 and 7 July 2005.

It is intended to lodge some of the collected specimens at the NHM, the remainder being kept in the author’s reference collection. Andrew Halstead has indicated that he intends to donate two females to the BENHS collection at Dinton Pastures.

Determination

The host plant list in Lacourt (1999) includes P. leucopus as the only European species of the genus found exclusively on Tilia. Application of the Pristiphora key in Zhelochovtsev (1988), treated there as a sub-genus of Nematus, enabled determina- tion of the British females as P. leucopus. The key characters are the extent of white on the hind femora, the sub-bifid tarsal claws and the larval host plant.

Revised key

This is a modification to Benson’s 1958 key to females in the B Group of Pristiphora. The revision begins by replacing Benson’s couplet 6 on p. 159 with the following and adding new couplets 6a and 6b:

6 (3) Hind femur mainly black 6a

Hind femur mainly yellowish white ,6b

6a (6) Hind femur mainly black, sometimes with extreme base and apex white, but

never more than 10% of total surface area white 7

Hind femur with extensive area of black but with a minimum of 30% white,

sometimes more 6b

6b Tarsal claws with a large sub-apical tooth, almost bifid. Some individuals emerging in May, after overwintering as prepupal larvae, show more extensive black on the hind femora than those emerging in the summer

generations which have mainly white femora P. leucopus Hellen

Larva on Tilia

Tarsal claws with inner tooth either absent or tiny 11

This key has been simplified to enable separation of P. leucopus, some of the other characters included by Benson would need to be reinstated in order to identify the species later in the key. The combination of white on the femora, sub-bifid tarsal claws and the use of Tilia as a larval hostplant precludes confusion with species which are superficially similar.

The male key is revised as follows, replacing Benson’s couplet 3 on p. 164 with the

following and adding a new couplet 3a:

3 (2) Hind femur and tibia reddish-yellow or yellowish-white 3a

Hind femur mainly black and tibia mainly brownish-white 6

BR. J. ENT. NAT. HIST., 19: 2006

5

3a Hind femur and tibia mainly reddish-yellow 4

Hind femur and tibia mainly yellow-white, claws with large sub-apical inner

tooth, penis valve (Fig. 4 in this paper) P. leucopus

It is not known whether British males emerging in May have black on the femora.

Acknowledgements

The author is grateful for the help of the following people in researching and compiling this paper. Neil Springate at the Natural History Museum, London for arranging the loan of specimens and searching for references. Beatrice Gillam, Andrew Halstead, Andrew Liston and Adam Wright for their extensive comments on early drafts. Andrew Halstead also contributed further by the loan of specimens taken by him at Wisley. John Badmin for sending me larvae for rearing which he collected in Norfolk.

References

Benson, R.B. 1958. Handbooks for the Identification of British Insects. Vol. VI Part 2(c).

Hymenoptera, Symphyta. Royal Entomological Society of London.

Lacourt, J. 1999. Repertoire des Tenthredinidae ouest-palearctiques. (Hymenoptera, Symphyta) . Societe Entomologique de France, Paris.

Lindqvist, E. 1969. Blattwespen-Studien (Hymenoptera, Symphyta). Notulae Entomologicae. XLIX. pp. 38-48.

Zhelochovtsev, A.N. 1988. Pereponchatokrylye Shestaja Tschast. Oprodelitel nasekomych evropeskoy tshasti SSSR. 3 (Hymenoptera) , 6: 268 pp. Leningrad. (In Russian). (English translation, 1994. Keys to the Insects of the European Part of the USSR. Vol. Ill Hymenoptera, Part VI Symphyta. E.J. Brill, Leiden, New York & Koln.)

SHORT COMMUNICATION

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), the multi-coloured Asian

ladybird, an etymological note. There have been several recent publications in 2005 {e.g. Roy et al., Ecology of the harlequin ladybird a new invasive species. British Wildlife, 16: 403^-07; Ware et al. The harlequin ladybird arrives in Britain: a threat to our native species? Bull. amat. Ent. Soc., 64: 175-186), and many articles in the media, on the arrival in the British Isles of the coccinellid Harmonia axyridis. These provide what seem to be unnecessarily alarmist accounts of possible detrimental effects of this species on our existing coccinellid populations as well as on the populations of a number of other insects. In producing these accounts of H. axyridis , which is known throughout most of the English speaking world by the common name of multi-coloured Asian ladybird, has been re-christened the Harlequin ladybird. The reason for this change can only be guessed at, but the new name has no doubt been chosen to reflect the wide range of colour varieties found in this species, reminding one of a harlequin clown. There is, however, a probably unintentional significance to this name. Harlequin derives from the Old French Herlequin or Hellequin, the leader of the Wild Host or troop of demon horsemen ( The Shorter Oxford English Dictionary on Historical Principles. 3rd edition, revised 1973, Clarendon Press, Oxford). Given the predicted results of the spread of this species it will be interesting to see whether its real attributes will be those of the Herlequin or those of a parti-coloured bespangled clown taking advantage of a vacant urban niche. - John Muggleton, 1 7 Chantry Road, Wilton, Salisbury, SP2 0LT

6

BR. J. ENT. NAT. HIST., 19: 2006

REVIEW

The Empidoidea (Diptera) of Fennoscandia and Denmark. IV Genus Hilar a. Fauna

Entomologica Scandinavica 40 by M. Chvala, (2005). 233 pp. Brill, Leiden, Boston. ISBN 90-04-14799-3. Price: 99€

This landmark publication is the fourth in the FES series covering the Empidoidea of Fennoscandia, the previous volumes having dealt with Hybotidae, Microphoridae, Atelestidae and the genus Empis of the Empididae. This volume deals with the difficult empidid genus Hilara including sections on adult morphology, classification, life history and zoogeography with backing maps and tabular summaries. The main ‘meat’ however is the key and accompanying descriptions which provide an authoritative treatment of the 90 species of Hilara recorded from the region.

In Britain, correct determination of the empidid genus Hilara has been feasible (though thwart with difficulties for the unwary) thanks to J. E. Collin’s 1961 revision (British Flies VI, Empididae, CUP). However, understanding of the continental fauna has suffered much from taxonomic and nomenclatural uncertainty which this revisionary work has gone a long way to resolving, at least in northern Europe. Fennoscandian dipterists will welcome this volume with open arms but there is however, much to commend this book to British dipterists too.

Firstly all the British species are included, as well as some of those which might yet be found here. Whilst Collin's key is serviceable, Chvala’s studies have unearthed fresh diagnostic characters which will inspire confidence as users navigate their way through the 151 couplets. The problems of keying out wet-preserved material remain as dusting characters (usually hard to see in alcohol) still feature in Chvala’s key, however, the abundance of additional morphological characters should provide adequate compensation. Individual species accounts are full and detailed and it is satisfying to note the very necessary provision of illustrations of the male genitalia and basitarsi for all species together with supportive figures of other characters as needed. Each account is very usefully rounded off with short summaries of distribution and biology including ecology and epigamic behaviour.

It is difficult to find fault with this book. I would have liked to have seen a systematic checklist as a separate table and I did find a reference quoted in the text which was not present in the Literature section. But these are very minor issues. The price is another matter and is guaranteed to induce a wave of apoplectic derision in many potential buyers whose personal or institutional funds won’t stretch that far. This is an excellent and very useful book which deserves to be on the shelves of every British dipterist interested in the Empididae. It is a great pity that the economics of the marketplace will most likely prevent that from happening.

Adrian Plant

Correction

BARRINGTON, R.D.G. & WHITE, M.C. The first record of multiple allelomorphism in a British butterfly: Coenonympha tullia (Muller) ssp. polydama (Haworth) (Lepidoptera: Satyridae). British Journal of Entomology and Natural History 18: 253-258.

Contrary to the impression given by Barrington & White in their recent paper, Butterfly Conservation did not issue a licence for the release of excess breeding stock of the Large Heath butterfly. Butterfly Conservation is a registered UK charity and non-governmental organisation with no powers to issue licences. Permission to take or release butterflies should be sought from landowners or statutory bodies e.g. English Nature, where appropriate.

BR. J. ENT. NAT. HIST., 19: 2006

7

THE MODERN BEE AND WASP ASSEMBLAGES (HYMENOPTERA: ACULEATA) OF WARWICKSHIRE’S CALCAREOUS QUARRIES AND SPOILHEAPS, AND THE CONSERVATION ISSUES FACING THEM

Steven J. Falk

Warwickshire Museum, Market Place, Warwick CV34 4SA stevenfalk@warwickshire.gov.uk

Abstract

A comparative review of the modern bee and wasp assemblages (Hymenoptera: Aculeata) of fourteen calcareous (mudstone, limestone and ironstone) quarries and spoilheaps in Warwickshire is given. The sites were assessed in terms of species diversity, presence of rare species and the quality of various habitat-related assemblages such as calcicoles. 186 species were recorded in total during the main study period from 1990-2002, and the best site (the Bishops Bowl-Bishops Hill Complex) produced a list of 128 species. Notes on some of the scarcer species are provided and discussion of the main conservation issues that affect bees and wasps at these sites. The danger of placing too much emphasis on Species Quality Indices is highlighted.

Introduction

Geologically speaking, Watsonian Warwickshire is a county of two halves. The north and west is largely underlain by pre-Jurassic mudstone, sandstone and igneous rocks overlain by much younger glacial drift that mostly give rise to base-poor soils. These soils once supported scattered heathland within a setting that remained heavily wooded until relatively late, representing the legendary "Forest of Arden’ which is now classified as "Ancient Arden’ by the Warwickshire Landscape Guidelines (Warwickshire County Council, 1993 parts 13) and forming part of the Midlands Plateau Natural Area. Today this part of Warwickshire has lost most of its heathland and bog, but still supports a variety of acidophilous or calcifugous plants and insects which occur predominantly or exclusively within this part of the county.

By contrast, the south and east is largely underlain by mudstone and limestones of Jurassic age, with a scattered drift cover. The Jurassic succession includes a narrow ironstone unit (the Marlstone) that forms some of the escarpments along the northern edge of the Cotswolds such as Edge Hill, plus several large outlying hills just north of the Cotswolds. The underlying geology of southern and eastern Warwickshire has given rise to more fertile, calcareous soils, mostly within the landscape zone known as the ‘Feldon’, which falls within the Midland Clay Pasture Natural Area. This had largely been cleared of its woodland by the time the Romans arrived, and it is likely that calcareous grasslands were widespread here over many centuries within a landscape of open field systems and grazed hillsides. During the twentieth century, most of the species-rich grassland here was lost. Today, perhaps as little as 50 ha of "unimproved’ ancient limestone grassland remains and most of it is fairly unimpressive in entomological terms.

Fortunately, the various underlying deposits of this area have been exposed through quarrying and the production of cuttings for the local road, railway and canal networks. The quarries in particular display widely varying characters and ages and now collectively support a diverse array of calcareous conditions with many

BR. J. ENT. NAT. HIST., 19: 2006

unusual plants and insects found nowhere else in the County. Calcareous spoilheaps are associated with some of these quarries and are sometimes more interesting than the quarries themselves. A few spoilheaps have also been produced following excavation of railway cuttings through limestone hills during the nineteenth century. The largest of these, Harbury Spoilbank, is included in this review.

Yet, in spite of the fact that some of the sites covered in this paper are amongst the most important calcareous sites in the entire British Midlands and some of the most interesting wildlife sites in Warwickshire, they experience a disproportionate level of threat. This includes the two sites that score highest for bees, wasps, butterflies and a range of other insects. These are not Sites of Special Scientific Interest (SSSIs), whilst lesser sites, often somewhat degraded through the effects of succession, are. It was this in mind that the author set out to collate detailed and reasonably comparative data for a variety of insect groups at fourteen of the potentially best calcareous sites in the county. The following paper concentrates on the bee and wasp information obtained, extending the published information on the aculeates of calcareous habitats furnished by publications such as Archer (1997) and Alexander (2003).

The geological deposits covered

(i) White Lias the oldest (late Triassic) and most northerly of the important calcareous deposits in Warwickshire, stretching obliquely across the county from the Stour Valley in the south to near Rugby in the east. The White Lias comprises up to 2 metres of hard, fine-grained limestone that was once a popular building stone obtained from a small number of shallow quarries. White Lias can give rise to strongly calcareous habitats with many unusual calcicoles and the clay component can lead to the formation of wetlands and seasonal water- logging.

(ii) Wilmcote Limestone -a fine-grained late Triassic -early Jurassic limestone historically quarried from a few sites west of Stratford and also giving rise to strongly calcareous conditions.

(iii) Blue Lias -an early Jurassic deposit underlying much of southern and eastern Warwickshire. It comprises alternating beds of mudstone and fine-grained muddy limestone which has been exploited by the cement industry, resulting in several very large quarries and cement works, one of which (Rugby Works) is still operated by the company RMC Group PLC. Blue Lias produces a strongly calcareous soil prone to intense drying out in dry summers, but also capable of producing some fine wetlands and seasonally flooded areas.

(iv) Marlstone-a hard, tawny-coloured ‘ironstone’ of early Jurassic age that forms the famous ‘Hornton Stone’ so characteristic of the architecture along the northern edge of the Cotswolds. Several large, shallow quarries exist in the Edge Hill district and can give rise to a mildly calcareous habitat that lacks the floristic diversity and large number of calcicolous plants typically associated with the Lias quarries. The ironstone was also used as a source of ferrous oxide in medieval times, being obtained from shallow quarries on various hill tops in the same general area. This has resulted in some very uneven landforms, most of which are grassed-over agriculturally improved sheep pasture not covered by this paper.

(v) Middle Jurassic limestone - quarried most recently at Cross Hands Quarry at the southern tip of the county. The Middle Jurassic succession includes oolitic building stones such as the Chipping Norton Limestones, chalkier, less hardy stone such as Clypeus Grit, and substantial layers of mudstone and sandstone.

BR. J. ENT. NAT. HIST., 19: 2006

9

The limestones give rise to a more distinctly calcareous habitat than Hornton Stone, with many of the unusual calcicoles associated with Lias sites, though they do not easily produce wetlands.

The survey sites

Fourteen sites are considered, plus a combined list for Sites 2 and 3, which are immediately adjacent to one another and best considered as a single ecological unit (though traditionally they have been treated separately). A brief description of each site is given, plus the dates of visits. Dates in brackets refer to relatively short visits or visits in sub-optimal weather.

1. Avon Hill Quarry (AH) SP416505 a medium-sized ironstone and sandstone quarry with patches of species-rich grassland, tall herb, wetland (with seasonal water bodies, swamp and carr) and extensive patches of ruderal habitat maintained by regular four-wheel drive meetings. Dates of visits:

14.111.1997, 15.viii.2001, 30.iii.2002, 19.iv.2002, l.vi.2002, 6.vii.2002, 2.ix.2002, 19.x. 2002.

2. Bishops Bowl (BB) SP385588 (Fig. l)-a large Blue Lias quarry system created by the cement industry with a complex mosaic of species-rich calcareous grasslands, tall herb, several permanent water bodies (supporting commercial fishing), assorted wetlands and dense scrub/secondary woodland. Dates of visits: 10. iv. 1992, 7.V.2001, (8.V.2001), ll.vi.2001, 23.vii.2001, 27.vii.2001, 29.iii.2002, 21.iv.2002, 8.vi.2002, (12.vii.2002), 12.viii.2002, 13.ix.2002.

3. Bishops Hill (BH) SP393585-the large spoilheap and former works directly east of the previous site, now supporting species-rich grassland, tall herb and ruderal habitats plus extensive scrub and secondary woodland and a single small quarry. The site was extensively bulldozed in the late 1980s but has become flower-rich since. Dates of visits: 19.vi.1993, 17. ix. 1993, 20.iv.1994, 31.V.1994, 9.iv.l995,

30.111.1997, 25.vii.1998, 8.V.2001, 25.V.2001, 30.viii.2001, 30.vi.2002, 18.X.2002. Bishops Bowl + Bishops Hill (BB + BH)-the combined lists of the previous two sites, which are best treated as a single ecological unit for evaluation and designation purposes.

4. Cross Hands Quarry (CH) SP269290 (both sides of A44 road) -a pair of

medium-sized Middle Jurassic limestone quarries which form the southern tip of Warwickshire, supporting species-rich calcareous grassland, scrub and ruderal habitat. The larger quarry at the east (which includes a small geological SSSI) has largely been land-filled since the mid 1990s and lost much of its original interest. Dates of visits: 19.iii. 1995, 4.iv.l995, 28. vi. 1995, 27. iv. 1996,

3 1 .viii. 1 996, (6.iv.2002), 5.V.2002, l.vii.2002, 21.ix.2002.

5. Gypsy Hall (Wilmcote) Quarry (GH) SP 152592 -a medium-sized Wilmcote Limestone quarry containing a small geological SSSI, with species-rich calcareous grassland and surrounding scrub. Dates of visits: 24.vii.1999, 5.iv.2002, 3. v. 2002, 17.vi.2002, 16.vii.2002.

6. Harbury Spoilbank (HS) SP385598-a fairly small 6.7 ha site supporting a

mixture of species-rich calcareous grassland and tall herbs, with areas of dense scrub. Formed on a long hillock of Blue Lias spoil resulting from construction of the adjacent Leamington Spa to Oxford Railway line through an adjacent cutting in the 1840s. Part of a larger SSSI (Harbury Railway Cutting SSSI) now managed by Warwickshire Wildlife Trust. Dates of visits: 1 0.iv. 1 992,

1 3.vii. 1 997, 12.iv.2002, 12.V.2002, 12.vi.2002, 3.viii.2002, 14.viii.2002.

10

BR. J. ENT. NAT. HIST., 19: 2006

7. Lighthorne Quarry (LQ) SP324558-a small shallow White Lias quarry with some established calcareous grassland, sparsely-vegetated scree, several temporary pools and patches of grey willow. No formal management, but occasional sheep grazing. Dates of visits: 24. vi. 1999, 7.vii.l999, 14.viii.2001, 26.iii.2002, (5.iv.2002), 4.V.2002, l.ix.2002.

8. Napton Quarry (Na) SP455613-a large quarry system on the west side of Napton Hill consisting of a very old ironstone quarry at the top (containing a geological SSSI) and younger, but abandoned, brick works at the bottom. These collectively support much improved pasture, wetlands of various sorts (including pools, swamp, carr and spring-fed seepages), and ruderal/tall herb habitats. Dates of visits: 22.viii.1995, 5.iv.l996, 21. iv. 1996, 13. vi. 1996, 1 l.ix.1999, 7.iv.2002, 5.V.2002, 28.vi.2002, 24.vii.2002, 22.ix.2002.

9. Nelsons Quarry (Nel) SP443642 (Fig. 2) -a deep Blue Lias quarry with large spoilheaps supporting calcareous grassland, ruderal habitat, scrub, several water bodies with surrounding swamp and reedbeds, secondary woodland and a large seepage system. Part of a larger SSSI (Stockton Quarries and Cutting SSSI) also containing Stockton Cutting. Dates of visits: 1 2.vi. 1 994, 9.iv.l995, 13.iv.2002, 13. v. 2002, 13.vii.2002, 2 1 .viii.2002.

10. Newbold Quarry LNR (New) SP494769- another deep, water-filled Blue Lias quarry, in the suburbs of north Rugby. Extensively scrubbed over but with limited areas of species-rich calcareous grassland. A Local Nature Reserve managed by Warwickshire Wildlife Trust. Dates of visits: 23. iv. 1995, 16.iii.1997, 7.iv.2002, 2.vi.2002, 14.vii.2002, l.ix.2002, 18.X.2002.

1 1. Ratley Grange (Edge Hill) Quarry (RG) SP371470. A large Marlstone quarry on the top of Edge Hill, which was being actively worked during the survey period (in a pattern that permitted plentiful ruderal habitat to form), but currently lacking much of its original interest and likely to be developed. Dates of visits: 25. iv. 1996, 7.vii. 1 996, 27.viii.1996, 2 1 .viii. 1 997, 8.vi.2001, 4.V.2002.

12. Southam Quarry (So) SP421634. A very large series of quarries and spoilheaps, with exposures of White Lias and Blue Lias, still partially worked by RMC for cement and expanding. Very extensive species-rich calcareous grassland and ruderal habitats, scrub and limited wetlands. Dates of visits: 26.V.1998,

1 1. viii. 1999, 23.V.2001, 23.vi.2001, 10.viii.2001, 27.iii.2002, 24.iv.2002,

19. vi. 2002, 27.vii.2002, 1 1 .viii.2002.

13. Stockton Cutting (St) SP440649. A disused railway cutting plus adjacent Blue Lias quarries with remains of former works, supporting areas of species-rich calcareous grassland, tall herbs and extensive scrub and secondary woodland. Part of a larger SSSI (Stockton Quarries and Cutting SSSI) and a Local Nature Reserve managed by Warwickshire Wildlife Trust. Dates of visits: 27.vi.1990,

1 2. vii. 1990, 9.vi. 1 99 1 , 12.V.1996, 1 1 .viii. 1996, (25.V.2001), 29.iii.2002, ll.iv.2002, 19.V.2002, ll.viii.2002, 6.X.2002.

14. Ufton Fields SSSI (UF) SP381614-a 32 ha former White and Blue Lias quarry

back-filled to form ridges and furrows plus flatter areas. Now supporting a complex mosaic of species-rich grassland and tall herb, scrub, water bodies, swamp, carr and secondary woodland. Also a Warwickshire County Council Country Park and Warwickshire Wildlife Trust reserve. Dates of visits: l.v.1990, 23. v. 1990, 9.vi.l99L ll.iv.1992, 9.ix.l995, 15.iii.1997, (8.V.2001), 30.vii.2001, 14. viii. 2001, 26.iii.2002, 6.iv.2002, ll.v.2002, 23.vi.2002,

18. viii. 2002, 15.X.2002.

BR. J. ENT. NAT. HIST., 19: 2006

Figure 1. Bishops Bowl.

Figure 2. Nelsons Quarry.

12

BR. J. ENT. NAT. HIST., 19: 2006

Survey methodology

Surveying took place between 1990 and 2002 with a structured programme of visits in 2001 and 2002 to ensure that most sites received at least five visits extending from March/April to September/October. Some sites received considerably more visits than this due to circumstances (such as the preparation of impact assessments), though by the end of 2002 it had become difficult to extend the site lists much further for bees and wasps. A variety of survey techniques was employed, including visual surveillance of foraging and nesting habitats and careful sweeping of different habitats with a long-handled net, including patches of flowers, sparsely vegetated areas and sunlit foliage. Special attention was given to the flowers that supported the biggest foraging assemblages at particular times of year, which included:

Spring: grey willow Salix cinerea , blackthorn Prunus spinosa , hawthorns Crataegus spp., dandelions Taraxacum spp., daisy Beilis perennis , coltsfoot Tussilago farfara , ground-ivy Glechoma hederacea and dead nettles Lamium spp.

Early summer: mouse-ear hawkweed Pilosella officinarum , hawk’s-beards Crepis spp., birds-foot trefoils Lotus spp., kidney vetch Anthyllis vulneraria, vetches Vicia spp., hogweed Heracleum sphondylium, oxeye daisy Leucanthemum vulgare and bramble Rubus fruticosus agg.

Mid to late summer: thistles Cirsium and Carduus spp., ragworts Senecio spp., knapweeds Centaurea spp., wild parsnip Pastinaca sativa, wild carrot Daucus carota mignonettes Reseda spp., rosebay willowherb Chamerion august if olium, melilots Melilotus spp. and hawkweed ox-tongue Pier is hieracioides.

Late summer-early autumn: perennial sowthistle Sonchus arvensis, scentless mayweed Tripleurospermum inodorum , hawkbits Leontodon spp., angelica Angelica sylvestris and late flowers from the previous category.

Persistence and timing were important for producing good lists. The males of Lasioglossum xanthopus (Kirby), for example, peak in late September in Warwick- shire, a time when many hymenopterists have ceased recording for the year. Other species such as Andrena praecox (Scopoli) start foraging so early that they can have peaked by early April in early springs and are difficult to find by the end of this month. Finding species with very narrow foraging habits requires careful surveillance of specific flowers e.g. mignonettes for Hylaeus signatus Panzer and willows for Andrena apicata Smith, A. clarkella (Kirby) and A. praecox. For groups containing species that are indistinguishable in the field such as Sphecodes, Lasioglossum , Crossocerus and Pemphredon, reasonable-sized samples were obtained for critical checking under a microscope.

Three parameters were then used to assess the quality of the assemblages present:

(i) Species richness. This figure was the total number of bee and wasp species recorded per site during the survey period.

(ii) The presence of rare species. The rarity gradings for Red Data Book and Nationally Scarce species used for assessing this parameter were taken from Falk (1991). However, to allow for the fact that some of these gradings are now known to be misleading, an asterisk has been placed against the obviously mis- graded species in Appendix 2 and a bracketed re-grade suggestion that is more realistic given afterwards. Regional scarcity was assessed using information in the available national atlases published by the Bee, Wasp and Ant Recording Society -‘BWARS’ (Edwards, 1997, 1998; Edwards & Telfer, 2001, 2002); also personal data and other literature or correspondence to hand. A rarity score (RS) was evaluated for each site by assigning points to the various rarity

BR. J. ENT. NAT. HIST., 19: 2006

13

gradings as follows: 100 points to Red Data Book species, 50 points to Nationally Scarce species and 20 points to Regionally Scarce species (following Ball, 1986). The rarity score was also divided by the total number of species at a site to produce a Species Quality Index (SQI) which is stated to even out variable recording coverage across multiple sites. These calculations were made for each of the fourteen study sites (plus sites 2 and 3 combined), following adjustment for obvious misgradings.

(iii) The quality of certain habitat-associated assemblages. A variety of habitat- linked insect assemblages can be used to compare site quality within certain defined parameters. At the sites studied here, these included assemblages associated with open calcareous habitats, shaded calcareous habitats, calcareous wetlands and general wetlands. The first was the main one relevant to the bees and wasps and such species are flagged as calcicoles (Calc) in Table 1 and Appendix 2. A single wetland specific (Wetl) bee was also present at one site, but no bee or wasp species recorded were specialists of shaded calcareous habitats or calcareous wetlands (categories that are important for flies and some other groups). Across the insect fauna as a whole, an interesting assemblage can develop in association with snails at calcareous sites, so a category for snail- associated species was used (Sn), many of which also fall into the calcicole category. These are mostly predatory and parasitic flies and beetles, but include a small number of bees that nest in empty snail shells. The number of aerial nesters at each site i.e. those species nesting in dead wood, hollow stems or amongst foliage (e.g. Dolichovespula wasps), was also used as a further parameter specific to bees and wasps. The assignment of species to the calcicole category was based on information in national atlases, other literature and personal data. However, it should be noted that some species that act as calcicoles in Warwickshire, fail to do so in other parts of Britain. This includes Andrena flavipes Panzer, Odynerus melanocephalus (Gmelin in L.), Pseudospi- nolia neglecta (Schuckard), Priocnemis parvula Dahlbom, Sphecodes ferruginatus von Hagens and S. hyalinatus von Hagens. For Sphecodes hyalinatus it stems from the fact that the main Warwickshire host is the calcicolous Lasioglossum fulvicorne (Kirby) rather than the acidophilous L.fratellum (Perez) which acts as the main host in many other parts of Britain.

Results

Species richness

Good samples of aculeates were obtained from all fourteen study sites, to the extent that it started to prove difficult to extend the site lists further by the end of the survey period. The data were used to evaluate the quality of the assemblages present at each of the sites (plus sites 2 and 3 combined). The various scores that underpin this evaluation are presented in Table 1. Brief status notes on the scarce species are given in Appendix 1, together with a full species list for each site in Appendix 2.

186 species of bee and wasp (excluding Apis mellifera L.) were recorded with certainty from the fourteen sites. The Bishops Bowl- Bishops Hill Complex produced the longest list, of 128 species. Within the West Midlands Region ( sensu Herefordshire, Worcestershire, Warwickshire, Staffordshire Shropshire and the former West Midlands County), this is only surpassed by Highgate Common, Staffordshire which has recently produced a list of 130 species (S. Falk and

14

BR. J. ENT. NAT. HIST., 19: 2006

M. Archer, combined data). Southam Quarry produced a list of 1 12 species. None of the other sites exceeded 100, and the poorest (Lighthorne Quarry) only produced 51 species.

The presence of rare and scarce species

Some very significant records were obtained, including several new county records, substantial national range extensions and numerous records for Red Data Book, Nationally Scarce and Regionally Scarce species. All sites produced records of rare or scarce species, though the number of these, and the SQI value they produced, varied greatly (see Table 1), though the figures were loosely correlated with species richness.

Southam Quarry and the Bishops Bowl-Bishops Hill Complex supported particularly important assemblages of scarcer bees and wasps (24 and 27 species, respectively). Their rarity scores of 1010 and 970, respectively, are probably the highest values for any bee and wasp assemblages in Vice-county Warwickshire and compare well, for example, with some of the better heathlands in the West Midlands Region. The scores were considerably higher than that recently obtained for Sutton Park National Nature Reserve (470) though some way short of that for Highgate Common (1800). These two sites collectively produced records of four rare or scarce species unknown from any other sites in Warwickshire at the time of the survey: Andrena fulvago (Christ), A. proxima (Kirby), Hylaeus pectoralis Forster and Osmia aurulenta (Panzer). They also supported important populations of Bombus ruderatus (Fab.), Bombus humilis Illiger and Nomada ferruginata (see Appendix 1). The poorest site for scarce species was Newbold Quarry, with only a single Regionally Scarce species. But even relatively impoverished sites have the ability to support surprisingly rare species. Lighthorne Quarry, the smallest and least rich site, with only six scarce species, still produced a record of the RDB1 N. ferruginata. Napton Quarry produced the second highest SQI through the presence of N. ferruginata , six Nationally Scarce species and nine Regionally Scarce species within its fauna of only 70 species. But given that the site was clearly nowhere near as rich as the Bishops Bowl-Bishops Hill Complex or Southam Quarry, and had a much lower rarity score, this reveals the danger in placing too much emphasis on SQIs, an evaluation system that is often used unquestioningly in site evaluation and one that can obscure substantial real differences in site quality.

In total, nine species were unknown in Warwickshire from any other sites except those considered here at the time of the survey. In addition to the four listed above, these were Arachnospila minutula (Dahlbom), Caliadurgus fasciatellus (Spinola), Lasioglossum xanthopus (Kirby), Sphecodes niger von Hagens and S. rubicundus von Hagens (Appendix 1). The records of H. pectoralis and O. aurulenta are still the only ones the author is aware of in the West Midlands region and represent considerable extensions to their known national ranges. Andrena proxima has been discovered at Napton Quarry since the survey finished, but this site and Bishops Bowl are still the only two locations known within the region.

The quality of certain habitat-linked assemblages

Twenty-one calcicoles, 62 aerial nesters (excluding cleptoparasites), three snail- users and one wetland specialist were recorded. The numbers of these found at each site are summarised in Table 1. For calcicoles, Southam Quarry and the Bishops Bowl Bishops Hill Complex produced a substantially higher score than the other

BR. J. ENT. NAT. HIST., 19: 2006

15

Ufton Fields SSSI (UF)	o , — i O t — co o “cp On - — i o co OO ’— 1 CN d"
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BR. J. ENT. NAT. HIST., 19: 2006

sites (15 and 13 calcicoles, respectively), reflecting the extent and quality of the open calcareous habitats present. This compares to only two such species at Newbold Quarry. These sites also supported a much higher number of aerial nesters (35 and 33 such species, respectively) compared to just five such species at Gypsy Hall Quarry. Just these two parameters considered alone can suffice to reflect the quality of a calcareous site as they are functions of site size, habitat quality and habitat diversity, and a simultaneous high score in both parameters reflects all three attributes. Bishops Bowl produced the only record of a wetland aculeate, Hylaeus pectoralis, reflecting the presence of extensive reed beds. The numbers of calcicoles and aerial nesters were loosely correlated with species richness.

Discussion

Factors influencing species richness

(i) The extent of floristically diverse habitat. The Bishops Bowl-Bishops Hill Complex and Southam Quarry supported the largest expanses of floristically- rich limestone grassland and early successional habitat, with very extensive stands of many key forage plants such as spring-blossoming shrubs, birds-foot trefoils, kidney vetch, various umbellifers, thistles, ragworts, knapweeds, oxeye daisy and scentless mayweed. Species such as Bombus ruderatus , B. humilis and Osmia aurulenta appear to specifically require large areas of flower-rich habitat and depend heavily on plants like birds-foot trefoils. The poorest sites (those with under 70 species recorded) were either characterised by relatively small areas of flower-rich habitat (usually much reduced by scrub encroachment or excessive disturbance) or by a lack of floristic diversity or lack of certain key forage plants within the open habitats that were present. The latter occurred at the three sites featuring Ironstone (only mildly calcareous compared with Lias deposits and lacking many key calcicoles); also at sites where the ground conditions were too xeric for a rich plant community. Blue Lias clay for example can often hold very little water in a dry summer, and in the driest areas it can take several decades to develop a reasonable vegetation cover which may only contain the most drought tolerant species such as dog-rose Rosa canina agg., dwarf thistle Cirsium acaule and autumn gentian Gentianella amarella.

(ii) The quality of habitat mosaics and adjacent habitats. Even where sufficiently large areas of flower-rich, calcareous habitats were present, the variety of other habitats and successional stages in and around a quarry was highly influential on richness. Woodland, scrub, and tall herbs can provide important foraging habitats and provide nest sites for a good variety of aerial nesters. Habitats close to a quarry system such as flowery road verges, fields margins, hedges, fallow fields, disused railways and canal banks, even gardens can also boost the aculeate diversity of a quarry, and often support large stands of critical forage plants (especially thistles, hogweed and scentless mayweed) that may be scarce or absent within the quarry itself. It is important to be aware of the landscape around a quarry site when trying to understand the factors influencing the bee and wasp fauna, and to be aware of the likelihood that in some aculeate species local populations are operating at a landscape level across a cluster of suitable sites.

(iii) Geology and site history. It is thought that geology can influence the precise composition of bee and wasp assemblages, though it is difficult to separate the affects of geology from the other variables at play, particularly the history of a

BR. J. ENT. NAT. HIST., 19: 2006

17

site and the pattern of operations associated with its active phase. The large Blue Lias quarries studied appear to have the greatest potential for bees and wasps, possibly because of the piecemeal pattern of operations usually associated with the cement industry. This tends to produce large spoilheaps and other areas that are allowed to re-vegetate naturally at different times and rates, and to create new quarries adjacent to older, abandoned ones, resulting in complex habitat mosaics. Clay-rich overburden was often mounded up and can produce different conditions to pure Lias deposits. The inability of ironstone quarries to support certain key calcicolous plants has already been mentioned. Uneven quarry floors can result in a complex mosaic of wetland conditions ranging from permanent deep water through to seasonally-flooded inundation marsh and damp ditches and hollows. This can promote the abundance of valuable forage plants such as grey willow, angelica and marsh thistle Cirsium palustre, which prefer damp soils. As a rule, longer-abandoned sites tended to exhibit more advanced levels of succession such as scrub encroachment or the development of coarse grassland and have often lost much of their early successional habitats and the aculeates that require these. Rabbit activity and informal human disturbance can sometimes slow down the rate of succession or completely arrest it locally.

(iv) Topography and altitude. A number of the sites were located on fairly high ridges or hills e.g. Cross Hands Quarry, Napton Quarry and Ratley Grange Quarry. These tended to be more exposed to prevailing winds and on a given day (especially a windy one) were often somewhat cooler than nearby sites in less exposed locations. This appeared to reduce species diversity. Newbold Quarry suffered from the fact that much of its species-rich limestone grassland was on north-facing slopes or overshadowed by wooded slopes to the south. This reduced the extent of warm habitat available for thermophilic insects such as bees and wasps.

Factors influencing the number of rare and scarce species and key habitat-linked assemblages

These appear to be much the same as those influencing species richness, and the presence of scarcer species seemed strongly influenced by species richness. Larger sites with extensive, flowery calcareous habitat within a larger habitat mosaic had greater capacity for supporting the key forage plants and ideal nesting sites required by scarcer species and calcicoles. But for some individual rare species and calcicoles, it may have been the abundance of just a small number of flower species that supported their presence. Andrena proxima owed its occurrence at Bishops Bowl to a large patch of ground-elder Aegopodium podagraria. At Southam Quarry, Botnbus humilis, B. ruderatus and Osmia aurulenta were heavily reliant on the very extensive birds-foot trefoils and kidney vetch there. The rare cleptoparasite Nomada ferruginata was indirectly dependent on the willows that feed its sole host Andrena praecox. Hylaeus signatus was entirely dependent on wild mignonette Reseda lutea and weld R. luteola.

Evidence of losses and gains in the bee and wasp fauna

The low level of insect recording at most of the study sites prior to the 1980s hinders our knowledge of losses. The only certain loss is that of Bombus sylvarum (L.), which was present at Ufton Fields as late as 1965 (the last record for

18

BR. J. ENT. NAT. HIST., 19: 2006

Warwickshire). However, extensive infilling and landscaping of Cross Hands and Ratley Grange Quarries have almost certainly resulted in losses here over recent years, including half of the known Warwickshire sites for Lasioglossum xanthopus. Excessive scrub encroachment has probably reduced the diversity of at least four further sites (see Vegetation succession below). Rather more evidence is available for recent gains, and this mostly relates to species known to be expanding their ranges nationally. These include Andrena flavipes, Crossocerus distinguendus (A. Morawitz), Dolichovespula media (Retzius), D. saxonica (Fab.), Ectemnius rubicola (Dufour & Perris), Lasioglossum malachurum , Microdynerus exilis and Philanthus triangulum (Fab.), and possibly also Didineis lunicornis (Fab.), Andrena proximo, Hylaeus cornutus Curtis, Hylaeus pectoralis and Sphecodes niger. Climatic factors appear to underlie these expansions, and the sites considered here appear to represent important stepping stones for such expanding species (the intervening countryside providing few opportunities). These records indicate the dispersal abilities of such species, which is usually poorly documented.

The relative national importance of the study sites for bees and wasps

The level of species richness and number of rare and scarce species encountered across the fourteen study sites was comparable to that associated with lowland heathland in the West Midlands (S. Falk data). In the West Midlands Region, any modern list exceeding 100 species can be regarded as extremely good, though heathland sites in counties such as Surrey and chalk heath sites in the East Anglian Brecks by comparison still occasionally produce lists of 200 or more species (S. Falk, D. Baldock & J. Field data). Rather surprisingly, the best calcareous sites on the East Sussex downs, including a number of National Nature Reserves and SSSIs, do not appear to be much richer than the best sites covered here (S. Falk data). This may reflect the less complex topography and habitat mosaics associated with many ancient downland sites compared with large quarries. Based on such comparisons, sites such as the Bishops Bowl Bishops Hill Complex and Southam Quarry ought to be viewed as nationally significant. Southam Quarry is also noteworthy in that it produced records for 14 Bombus species (9 non-parasitic ones and 5 ‘cuckoo’ species). It appears to be the richest bumblebee site in the British Midlands today.

General factors affecting the bee and wasp fauna at these sites

( i ) Vegetation succession

The intensity of operations at many quarries during their peak of productivity can limit their entomological and botanical interest. Diversity increases as soon as quarries are partially or completely abandoned and flower-rich vegetation starts to establish. The optimal condition for bees and wasps will tend to occur once a range of floristically diverse conditions, representing a variety of successional stages, has developed. This may take 10-30 years after abandonment, depending on the size of a site and factors such as rabbit levels, hydrology, soil chemistry, and physical disturbance. Flowery early successional stages with plants such as oxeye daisy, hawk’s-beards, birds-foot trefoils and kidney vetch are vital for many aculeates, but areas of scrub, bramble and tall herb can also provide valuable foraging habitat and a source of nesting sites for aerial nesters. But, without further management or disturbance, excessive encroachment of such habitats or floristically poor grassland

BR. J. ENT. NAT. HIST., 19: 2006

.9

such as dense wood false-brome Brachypodium sylvaticum can swamp out valuable early successional stages, reducing the diversity of conditions and, in consequence, reduce the variety of bees and wasps. A number of the study sites have deteriorated in the past through the effects of succession, notably Ufton Fields SSSI, Stockton Quarries and Cutting SSSI (containing Stockton Cutting and Nelsons Quarry), Harbury Railway Cutting SSSI and Newbold Quarry LNR. SSSI notification made little difference. Most of these sites are now subject to active scrub control, but have lost a number of calcicolous insects as a result of their recent history, including butterflies such as the small blue Cupido minimus (Fuessley). It is presumed certain bees and wasps have been lost too, including some of the species currently confined to the Bishops Bowl-Bishops Hill Complex and Southam Quarry, which currently support the conditions once associated with these other sites.

( ii ) Quarry infilling and landscaping

Two sites have been subject to partial infilling and landscaping, Cross Hands Quarry and Ratley Grange Quarry. Currently, it is likely that most of the scarcer bees and wasps recorded there during this study have been lost. The re-establishment of flower-rich vegetation may draw some of these species back, though it awaits to be seen whether the restored sites will regain high quality habitat. However, there is proven potential for land-filled and restored quarries and spoilheaps to regain high entomological value within a decade or two if capped with low fertility sub-soil sourced from other parts of the quarries and allowed to re-vegetate naturally over a sufficiently large area. Bishops Hill, for example, was subject to major re-profiling in the late 1980s, which was viewed as highly damaging to the ecology at the time, but the new ground had already acquired floristically diverse conditions and strong populations of many scarce bees and wasps by the mid-1990s. The most valuable part of Southam Quarry today is a re-vegetated mound of clay that was bare only a couple of decades ago. Where landscaping for nature conservation is taking place, it is essential that the introduction of top soil is avoided and that natural plant regeneration directly from the indigenous seed bank is encouraged. It is also important to produce plenty of south-facing slopes, including banks and some vertical faces.

(Hi) Development

During the time of writing, planning applications for development proposals that could substantially impact on site quality had been submitted for Bishops Hill, Bishops Bowl, Ratley Grange Quarry and the lower part of Napton Quarry. This is not surprising given that all the sites covered here fall within a developer’s concept of ’brown-field’ land which is still unquestioningly viewed as a more acceptable location for development than ‘green-field’ land (even though the latter is predominantly ecologically impoverished farmland in Warwickshire). Some of the impact assessments associated with these proposals have been very deficient, either due to poor expertise on the part of the ecological consultants working for the applicants, or a lack of sufficient resources or time for bona-fide ecologists to carry out sufficiently detailed surveys, data searches and impact analysis. The development of Individual Species Impact Assessments by the author (Falk, 1998) was a direct response to a deficient impact assessment at one of the sites covered by this paper. Several of the abandoned quarries in the suburbs of Rugby have become surrounded by residential or industrial development over the past 20-40 years, which has left a legacy of rather

20

BR. J. ENT. NAT. HIST., 19: 2006

isolated quarries with little subsidiary habitat. The agricultural land surrounding other sites has become much more intensively farmed in the latter part of the twentieth century, reducing the amount of subsidiary foraging habitat and possibly exposing quarries to the effects of pesticide drift.

( iv ) Excessive disturbance

This is more of a perceived threat than a real one, and nearly all of the study sites would benefit from higher levels of periodic, piecemeal disturbance, especially where this keeps the encroachment of scrub and wood false-brome in check. Avon Hill Quarry currently permits regular meetings of four-wheeled drive vehicles, which follow set routes through an interesting configuration of humps and hollows. Some parts of the site receive intense pressure, but overall the effect is beneficial, resulting in a complex array of early successional stages with good patches of birds-foot trefoils, mouse-ear hawkweed, melilots and clovers. Even where examples of large- scale disturbance have taken place, calcareous substrates have generally shown a remarkable ability to regain floristically diverse conditions with plentiful calcicoles once disturbance ceases.

(v) Site designation

Four of the fourteen sites constitute biological SSSIs: Harbury Spoilbank, Stockton Cutting and the nearby Nelsons Quarry (forming parts of one larger SSSI) and Ufton Fields. This affords legal protection from damaging operations, though, as noted above, it has not protected them from the insidious effects of vegetation succession in the past. Their aculeate faunas are now considerably less diverse then those of the Bishops Hill-Bishop Bowl Complex and Southam Quarry, with fewer rare species. Newbold Quarry is a Focal Nature Reserve and Ufton Fields is a Warwickshire County Council Country Park. These designations afford substantial protection as they are generally applied to land that is considered out of bounds to development proposals and under sympathetic ownership. Few of the remaining sites have yet to be formally designated as Second-tier Wildlife Sites (locally termed ‘SINCS’ -Sites of Importance for Nature Conservation), though most are flagged as ‘provisional SINCs’ or ‘Ecosites’ to alert local planning authorities of the need to treat planning proposals affecting them with appropriate vigilance. The Bishops Bowl-Bishops Hill Complex was designated as a provisional SINC during the preparation of this paper, largely on the basis of its entomological value and it is expected that others will follow. Entomological site data can be a powerful tool for designation where it is sufficiently detailed, comparative and interpreted.

( vi) Site management and monitoring

Three of the four SSSIs have been managed by Warwickshire Wildlife Trust (WWT) over a number of decades, with financial support through English Nature’s Reserves Enhancement Scheme. Newbold Quarry FNR is also a WWT Reserve. Ironically, it is these four sites that experienced the greatest past losses of flower-rich habitat through scrub encroachment. The substantial resources required for effective scrub control have been badly under-estimated in the past, and the ecology of many of the scarcer insects was poorly appreciated, resulting in a gradual loss of the extensive early successional stages required by many such species. But now that these issues are more fully appreciated, substantial efforts are being made to restore these

BR. J. ENT. NAT. HIST., 19: 2006

21

sites to their former value and WWT is better placed to achieve this than any other organisation in the county. However, sites including the Bishops Bowl-Bishops Hill Complex, Avon Hill Quarry, Lighthorne Quarry and Gypsy Hall are largely unmanaged and reliant upon rabbit grazing, physical disturbance unrelated to nature conservation and soil characteristics to keep succession in check. As such they must be considered vulnerable.

( vii ) Local Biodiversity Action Plans (LBAPs)

The bee and wasp fauna of Warwickshire’s calcareous sites is catered for in three habitat action plans in the Warwickshire, Coventry and Solihull LBAP: Quarries and Gravel Pits, Lowland Calcareous Grassland and Disused Industrial and Railway Land. These contain a variety of targets for habitat management, restoration, creation and designation. Species action plans have also been produced for Nomada ferruginata and jointly for Bombus humilis and B. ruderatus, three species featured in the National Biodiversity Action Plan (Anon, 1999). The various habitat and species action plans for Warwickshire can be viewed on the web at: www. Warwickshire. gov.uk/biodiversity.

Conclusions

Warwickshire’s calcareous quarries and spoilheaps contribute substantially to the biodiversity of bees and wasps in Britain, by allowing many species new opportunities for range expansion, by supporting a good number of scarce species, and by producing some very valuable calcicolous assemblages. The level of interest may be considerably greater than the ancient calcareous grasslands that preceded their existence (which are likely to have been less physically diverse). But threats to these assemblages are manifold and do not necessarily cease following protective designation, especially if insufficient management is taking place. Assessing the quality of sites should ideally examine the quality of habitat-associated assemblages in addition to noting species richness and the presence of scarcer species, but great caution is urged with the use of SQIs. The Southam Quarry and Bishops Bowl Bishops Hill Complex both require notification as SSSIs to help preserve their nationally important but vulnerable assemblages.

Acknowledgements

The author is grateful to the various site owners for permission to survey and for information relating to the histories of the sites. Dr J. Radley (Keeper of Geology, Warwickshire Museum) kindly assisted in the geological aspects of this paper. M. Edwards and G. Else provided some useful information on some of the scarcer species encountered.

References

Alexander, K. N. A. 2003. A review of the invertebrates associated with lowland calcareous grassland. English Nature Research Reports , Number 512. English Nature, Peterborough. Anon, 1999. Invertebrates. UK Biodiversity Group Tranche 2 Action Plans , Volume IV. English Nature, Peterborough.

BR. J. ENT. NAT. HIST., 19: 2006

22

Archer, M. E. 1997. The aculeate wasps and bees (Hymenoptera: Aculeata) of two calcareous localities in Watsonian Yorkshire: Burton Leonard Lime Quarries and Cave Wold. Naturalist 122: 45-52.

Ball, S. G. 1986. Terrestrial and Lreshwater Invertebrates with Red Data Book, Notable or Habitat Indicator Status. Invertebrate Site Register, 66 (Contract Reports, 637). Nature Conservancy Council, Peterborough.

Edwards, R. (ed.) 1997. Provisional atlas of the aculeate Hymenoptera of Britain and Ireland. Part 1. Bees, Wasps and Ants Recording Society. Huntingdon: Biological Records Centre.

Edwards, R. (ed.) 1998. Provisional atlas of the aculeate Hymenoptera of Britain and Ireland. Part 2. Bees, Wasps and Ants Recording Society. Huntingdon: Biological Records Centre.

Edwards, R. & Telfer, M. G. (eds.) 2001. Provisional atlas of the aculeate Hymenoptera of Britain and Ireland. Part 3. Bees, Wasps and Ants Recording Society. Huntingdon: Biological Records Centre.

Edwards, R. & Telfer, M. G. (eds.) 2002. Provisional atlas of the aculeate Hymenoptera of Britain and Ireland. Part 4. Bees, Wasps and Ants Recording Society. Huntingdon: Biological Records Centre.

Falk, S. J. 1991. A review of the scarce and threatened bees, wasps and ants of Great Britain. Research and Survey in Nature Conservation, 35. Nature Conservancy Council, Peterborough.

Falk, S. J. 1998. Individual Species Impact Assessments: a standardised technique for describing the impact of development proposals on critical invertebrate species. British Journal of Entomology and Natural History 11: 19-29.

Warwickshire County Council 1993. Warwickshire Landscape Guidelines (3 parts). Warwick- shire County Council Planning & Transport Department, Warwick.

Appendix 1 . Notes on some of the scarcer species

Andrena bucephala Stephens and Nomada hirtipes Perez — A. bucephala shows strong calcicolous tendencies in Warwickshire and the dozen or so sites fall into three categories: re-vegetated quarries and spoilheaps, railway cuttings, and south-facing Cotswoldian hillsides with patches of scrub. Foraging mostly occurs on hawthorn and field maple Acer campestre during May, and males can form conspicuous swarms around such blossoms. The communal nests (several females sharing a common nest entrance) are often associated with rabbit burrows within a scrub- grassland mosaic. Five of the strongest A. bucephala colonies support N. hirtipes, the nomad bees often revealing the precise location of the host’s nest. Both species are very rare abroad (G. Else, personal communication), making all British colonies highly significant.

Andrena dorsata (Kirby) -noteworthy for its rarity in Warwickshire -it is a frequent species in many parts of southern Britain, preferring heathland and coastal districts. The Avon Hill record (a single male on grey willow catkins) is one of only two modern ones for the county and its presence here may have been influenced by flowering gorse thickets (a major foraging habitat for first generation females at many sites) on nearby hills.

Andrena flavipes- almost certainly a recent colonist in Warwickshire, where first recorded in 1999 on agricultural land. All but two of the subsequent records fall within the sites covered here and suggest that calcareous quarries provide the most ideal habitat for it in Warwickshire.

Andrena fulvago- the Southam record (23 May 2001) is the only modern one for Warwickshire (1950s H.W. Daltry material from ‘Rugby’ exists at Coventry Museum). Only a single female was encountered, close to a patch of Crepis, a likely forage plant.

BR. J. ENT. NAT. HIST., 19: 2006

23

Andrena praecox- not considered especially scarce nationally. However, the Warwickshire populations are unusual in showing strong calcicolous tendency, preferring to nest in bare or sparsely-vegetated Lias clay at sites with plentiful willows. At the sites considered here, foraging was observed on grey willow, goat willow Salix caprea and (at the end of its flight period) white willow S. alba. The other two willow-requiring mining bees, A apicata Smith and A. clarkella (Kirby) lack strong calcicolous tendencies, indeed A. clarkella favours sandy, acidic sites in districts away from limestone and the populations encountered during this survey were very weak.

Andrena proxima-a strong population was discovered at Bishops Bowl, representing a considerable extension to its known range in Britain. Males were numerous on 7 May 2001 swarming over paths and visiting daisy flowers. Females were recorded on 1 1 June 2001 and 8 June 2002 mainly foraging on a large patch of ground elder, and hogweed to a lesser extent. It is suspected that this is another species increasing in range and frequency within southern Britain (S. Falk data) and it has been recorded at Napton Quarry in 2005 subsequent to the survey.

Arachnospda minutula- Bishops Hill and Southam Quarry support the only populations known in Warwickshire.

Bombus humilis- the Southam Quarry population is the stronger of the two modern ones known in Warwickshire (the other population may already have been lost since 1995). At Southam, a queen was recorded foraging on Lotus spp. on the relatively late date of 23 May 2001. Workers were also observed visiting Lotus and red bartsia Odontites verna in July and August. Compared with the closely-related B. pascuorum (Scopoli), B. humilis seems to have a much shorter foraging season involving fewer forage species.

Bombus ruderatus -at Southam, a queen was found foraging on Anthyllis flowers on 23 May 2001 and presumed workers (which are very difficult to distinguish from those of B. hortorum (L.) but often include fully melanic individuals) were foraging almost exclusively on Lotus until August. Like B. humilis , a relatively short foraging period and limited foraging scope were noted, compared to the closely-related B. hortorum. This BAP Priority Species seems to be exhibiting a major expansion in south Warwickshire at the time of writing (S. Falk data), which is rather surprising given that it has been traditionally regarded as one of the most seriously declined British bumblebees.

Caliadurgus fasciatellus - the closely approximated Southam Quarry and Stockton Cutting support the only populations known in Warwickshire, which are associated with sparsely-vegetated Blue Lias clay.

Crossocerus distinguendus - seemingly another recent colonist, first discovered in Warwickshire in 1998, since when it has been found at six sites of rather different character including one of the sites covered here. Cross Hands Quarry.

Didineis lunicornis only a single record from the present study (Ufton Fields), plus a further record for Newbold Quarry (R. Wright, personal communication), though many of the sites appear to provide ideal habitat (sparsely-vegetated clay with desiccation cracks). Its other four Warwickshire sites include sheep or rabbit grazed hillsides and railway cuttings.

Dipogon variegatus (L.)-the Avon Hill record is only the second for Warwick- shire. It was first recorded in July 1998 from a post-industrial site in Coventry, which has since been developed.

Ectemnius sexcinctus (Fab.) the Cross Hands and Southam records are the third and fourth for the county.

24

BR. J. ENT. NAT. HIST., 19: 2006

Hylaeus cornutus- the Southam record is only the second for Warwickshire, and the locality of the previous record (a post-industrial site in Coventry) has now been developed. At both sites it was recorded on wild carrot flowers.

Hylaeus pectoralis-a. most unexpected record representing a substantial extension to its known range in Britain. The site involved, Bishop Bowl, supports the largest stands of Phragmites of all the sites surveyed. The wasp nests in dead stems of reeds and the old galls of Lipara flies on reeds.

Hylaeus signatus- the three sites covered here repeat a pattern seen elsewhere in the county of a species with an almost complete reliance on ‘brownfield’ land, in association with Reseda spp. However, the colonies at calcareous quarries are relatively weak compared with those associated with post-industrial sites in the Coventry region during the 1990s, possibly because of the greater frequency of Reseda-x'xch. habitat in the Coventry area during the 1990s (much of which has now been developed).

Lasioglossum malachurum - seemingly another recent colonist in Warwickshire where it was first recorded in 1999. Seven of its ten Warwickshire sites are calcareous quarries and females forage on a wide variety of flowers, but especially composites such as daisy, hawk-beards and oxe-eye daisy. Observed nesting colonies have been associated with hard clay ground.

Lasioglossum xanthopus- Warwickshire records are currently confined to the four calcareous quarries covered in this paper. Females have been observed foraging heavily from ragworts and kidney vetch in May and June and nesting in vertical banks. Males have been found on perennial sowthistle flowers in late September. No evidence of its scarce cleptoparasite Sphecodes spinulosus von Hagens could be found, though both species occur in a disused railway cutting just over the Oxfordshire border (C. O’Toole, pers. comm.). Populations at two of its four sites (Cross Hands and Ratley Grange Quarries) have almost certainly been lost since the records were made through recent landfilling and restoration operations.

Microdynerus exilis (Herrich-Schaffer)-the Southam record is the second for Warwickshire where it was first recorded in August 1998 (a post-industrial site in Coventry that has since been developed).

Nomada ferruginata- the Napton population, which was first discovered in April 1996, is a strong one with good numbers observed around a nesting colony of the host Andrena praecox that forages on the plentiful grey willow here. The Bishops Bowl and Lighthorne Heath records relate to singletons, and the presence of only a few grey willow bushes at Lighthorne suggests this site does not support a secure population of the host. National data (Edwards & Telfer, 2002) suggests N. ferruginata may be currently expanding in southern Britain, and does not support its continued grading as an RDB1 species.

Odynerus melanocephalus - only two further Warwickshire sites are known beyond the nine covered by this paper, and all sites are characterised by the presence of bare or sparsely vegetated clay slopes with plentiful black medick Medicago lupulina- the source of its prey, the larvae of Hypera weevils (pers. observ.). The short-funnelled nests have been observed on a few such slopes, and contrast strongly with the long- funnelled nests of O. spinipes that tend to occur in vertical sand or clay faces (often the sandy overburden at the top of a limestone quarry face). The majority of Warwickshire records for the Odynerus cleptoparasite Pseudo spinolia neglect a are associated with O. melanocephalus , though a strong population was formerly associated with O. spinipes at Ufton.

Osmia bicolor- this is one of our most strongly calcicolous bees, with only two other known Warwickshire sites beyond the twelve shown here. Females have mainly

BR. J. ENT. NAT. HIST., 19: 2006

25

been observed foraging from birds-foot trefoils and kidney vetch, though visits to ground ivy, hawthorn, crab apple Malus sylvestris, willows, dandelions, brambles, violets Viola spp. and cowslip Primula veris have also been noted and males will visit composites like mouse-ear hawkweed and hawks-beards. Another strongly calcicolous megachiline bee Hoplitis spinulosa (Kirby) occurs alongside O. bicolor at many of its sites but forages almost exclusively on composites. The suitability of sites probably depends on the combination of plentiful forage plants combined with an abundance of empty snail shells in warm, sheltered locations fully exposed to the sun (the nesting site).

Osmia aurulenta- the Bishops Hill and Southam Quarry populations are remarkable for their isolated location in the centre of England far from any other known colonies. This is typically a species of calcareous coastal dunes with a more limited presence inland, mainly on chalk grassland. Foraging has only been observed from birds-foot trefoils and kidney vetch and it seems to require large quantities of such flowers, combined with an abundance of empty snail shells in open locations fully exposed to the sun for nesting.

Philanthus triangulum- singletons recorded from two sites. A recent colonist in Warwickshire, first recorded in 1995, with about a dozen records since. The sites studied here do not appear to be sandy enough to support strong colonies.

Priocnemis agilis (Schuckard)- Bishops Bowl is one of six Warwickshire sites, all of which are calcareous grasslands including semi-improved calcarous pasture.

Priocnemis parvula Dahlbom strongly associated with heathland and other sandy habitats over much of its range. Its presence at six of the calcareous sites studied here is noteworthy, though its presence at several non-calcareous sites locally has kept it off the calcicole list.

Psenulus schencki (Tournier)-the Ufton record is the second for Warwickshire, where it was first recorded in 1999.

Sphecodes niger the Nelsons Quarry record is the first for Warwickshire and coincides with a particularly strong colony of the host Lasioglossum morio (Fab.). Another species expanding nationally (S. Falk data, M. Edwards, pers. comm.).

Sphecodes rubicundus- the Napton Quarry and Bishops Bowl records were the only ones for Warwickshire at the time of the study (two more sites have subsequently been discovered) and coincide with strong populations of its main host, Andrena labialis (Kirby), which requires plentiful legume flowers.

Appendix 2. A list of all bees and wasps recorded at the 14 sites between 1990 and 2002.

The Status column lists various Quality Indicators. This includes species graded as Nationally Threatened (RDB1, 2, 3 & K), Nationally Scarce (N), Regionally Scarce ‘sensu Vice-county Warwickshire’ (R), calcicolous (Calc) and wetland-associated (Wetl). An asterisk after a grade indicates a misleading national grading and is followed by a bracketed value indicating a more realistic grading. A species name followed by an asterisk relates to species unknown from any other site in Warwickshire beyond those shown. Abbreviated site names are listed in the section on

26

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Appendix 2. A list of all bees and wasps recorded at the 14 sites between 1990 and 2002.

The Status column lists various Quality Indicators. This includes species graded as Nationally Threatened (RDB1, 2, 3 & K), Nationally Scarce (N), Regionally Scarce ‘sensu Vice-county Warwickshire’ (R), calcicolous (Calc) and wetland-associated (Wetl). An asterisk after a grade indicates a misleading national grading and is followed by a bracketed value indicating a more realistic grading. A species name followed by an asterisk relates to species unknown from any other site in Warwickshire beyond those shown. Abbreviated site names are listed in the section on

ACULEATE HYMENOPTERA	Rarity/ Quality Status AH BB B	BB + 1H BH CH GH HS LQ N	(a Nel New RG So St UF
Chrysididae Chrysis angustu/a Schenck Chrysis impressa Schenck Onuilus puncticoUis (Mocsary) Pseiulonuilus auratus (L.) Pseudospinolia neglect a (Schuckard) Trichrysis cyanea (L.)	R, Calc /		, -
Sapygidae ( Monosapyga clavicornis L.) Sapyga quinquepunctata (Fab.)	N ,
Tiphiidae Tiphia minuta Vander Linden	N* .(no / /	' ' ' ' ' ■
MutiUidae Myrmosa atra Panzer	/
Pompilidae Agenioideus cinctellns (Spinola) Anoplius concinnus (Dahlbom) Anoplius nigerrimus (Scopoli) Arciclinospi/a anceps (Wesmael)	R R

Araclmospila minutula (Dahlbom)* Arachnospila spissa (Schiodte)	N, Calc
Caliadwgus fasciatellus (Spinola)* Dipogon subintermedius (Magretti)	R
Dipogon variegatus (L.)	R
Priocnemis agilis (Schuckard) Priocnemis exaltata (Fab.)	N, Calc
Priocnemis parvula Dahlbom Priocnemis perturbator (Harris) Vespidae Ancistrocerus gazella (Panzer) Ancistrocerus nigricornis (Curtis) Ancistrocerus parietinus (L.) Ancistrocerus parietum (L.) Ancistrocerus trifasciatus (Muller)	R
Dolicbovespula media (Retzius) Dolichovespula norwegica (Fab.)	N* (no status)
Dolicbovespula saxonica (Fab.) Dolichovespula sylvestris (Scopoli)	RDBK* (no status)
Gymnomerus laevipes (Schuckard)	R
Micr odynerus exilis (Herrich- Schaffer)	N
Odynerus melanocephalus (Gmelin in L.)	N, Calc
Odynerus spinipes (L.) Symmorphus bifasciatus (L.) Symmorphus gracilis (Brulle) Vespula germanica (Fab.) Vespula rufa (L.) Vespula vulgaris (L.)	R

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Rarity /

ACULEATE Quality BB +

HYMENOPTERA Status AH BB BH BH CH GH HS LQ Na Nel New RG So St UF

Argogorytes fargeii (Schuckard) N

Cerceris rybyensis (L.) R

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Crossocerus dimidiatus (Fab.)

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Crossocerus elongatulus (Vander Linden)

Crossocerus megacephalus (Rossi us)

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Brulle)

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Didineis lunicornis (Fab.) N, Calc

Diodonius luperus Schuckard R

Ectemnius cavifrons (Thomson)

Ectemnius continuus (Fab.)

Ectenuiius rubicola (Dufour & Perris) R

Ectemnius sexcinctus (Fab. ) N

Entomognathus brevis (Vander Linden)

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34

BR. J. ENT. NAT. HIST., 19: 2006

NEW COUNTY RECORDS OF HETEROPTERA (HEMIPTERA) FROM GLOUCESTERSHIRE

Keith N. A. Alexander

59 Sweetbrier Lane, Heavitree, Exeter EX 1 3 AQ

Abstract

Eighteen species of terrestrial Heteroptera have been added to the Gloucestershire (vice-counties 33 and 34) county list over the last ten years. These have arisen from each of the main biogeographical regions of the county, and are largely from semi- natural vegetation. This suggests that the additions largely represent under-recording rather than recent colonisation.

Introduction

The Gloucestershire county Heteroptera fauna has been fully reviewed (Alexander, 1995 & 1996) and recording has subsequently aimed to consolidate coverage of the whole land area of the county and to confirm the continued presence of species which have not been noted for many years. One of the consequences of this new recording effort has been the discovery of a number of previously overlooked species as well as some which may be new arrivals. The two categories are often difficult to discern, however, as many bugs appear to be highly mobile and there are a large number which appear to be expanding their UK ranges northwards and westwards. The assumption is made that species which are strongly associated with long-established semi-natural vegetation are likely to be overlooked long-term residents. The eighteen additions bring the county total of terrestrial Heteroptera to 318 species, an increase of 6% in ten years.

Heathland fauna

A good case is presented by the county’s heathland fauna. Heathland was once very widespread in the Forest of Dean but has become very fragmented and localised through plantation forestry. Only recently has Forest Enterprise initiated some heathland conservation projects.

The absence of the most widespread heathland specialists from the county list was one of the most noticeable gaps, and investigation rapidly demonstrated their presence. The seed bug Scolopostethus decoratus (Hahn) (Lygaeidae) and the plant bug Orthotylus ericetorum (Fallen) (Miridae) were found at: Merring Meend Reserve, Plumphill (VC 34; S061), 15.ix.1990; Wigpool Common (VC34; S061), lO.viii. 1996; and Poor’s Allotment (VC34; ST59), 1 5.viii. 1993; as well as other sites subsequently, and are clearly ubiquitous in the heathland fragments. The predatory shieldbug Rhacognathus punctatus (L.) appears to be much more localised and may need larger areas of habitat. Nymphs were present amongst heather foliage at Crabtree Hill (VC34; S061)-a major Forest Enterprise heathland restoration area- lO.viii. 2002, and two adults were also found amongst the dense mosses beneath the heather canopy. Interestingly, only O. ericetorum has been found on the limestone heaths of the Cotswold Hills: Cleeve Cloud (VC33; S092), common on heather, 5.ix.l991.

BR. J. ENT. NAT. HIST., 19: 2006

35

Severn floodplain

The other major part of the county which appears to have been neglected by past recorders is the wide floodplain of the River Severn, with its expanses of modern intensively managed farmland and expanding urban and industrial developments. There are still fragments of the floodplain marshes however and these still support a few relict species.

The most interesting find has been the shore bug Saida littoralis (L.) (Saldidae). This is mainly a northern species in Britain, but also occurs in a few southern coastal counties. The northern habitat is typically river and lake margins where the marginal area is silty and there is vegetation close by, but in the south it is found in brackish habitats (Southwood & Leston, 1959). Its discovery in the old grazing marsh reserve of Ashleworth Ham (YC34; S082), 4.viii.2002, with its relict brackish water plant species, is a significant addition to the county list. Although the area still floods during the winter months the brackish nature of the water must be fairly negligible.

Other wetland species have also been found along the floodplain. The debris living predatory plant bug Fieberocapsus flaveolus (Reuter) has been found in small numbers by beating collapsed Glyceria stems over a net at Long Pool, Deerhurst (VC33; S082), 3.ix.2000-this area is part of a larger scheme by the Gloucestershire Wildlife Trust (GWT) to recreate areas of the former Severn marshes by returning areas currently under intensive agricultural exploitation to poorly-drained grazing land which is subject to natural flooding from the adjacent river.

The saltmarsh plant bug Orthotylus moncreaffi (Douglas & Scott) has also drawn attention to the county’s small areas of saltmarsh developed around the mouth of the Severn: Beachley Point (VC34; ST59), 14.vii.1990. Trigonotylus caelestialium (Kirkaldy) is the most recent addition to the county list, having been swept from tall saltmarsh grassland along the estuary at Whitescourt, Awre (VC34; SO70), 1 l.viii.2004.

The floodplain also appears to be an important area for tree canopy bugs. Ashleworth Ham is again the only county site for one of these species, the aphid- feeding plant bug Pilophorus clavatus (L.), associated principally with the broad- leaved willows Salix cinerea and S. capraea (Southwood & Leston, 1959), although beaten from hawthorn here, 4.viii.2002. The similar P. perplexus Douglas & Scott has also been added to the county list recently, and is proving to be scattered along the margins of the floodplain. It is another ant mimic, and similarly feeds on aphids and other soft-bodied invertebrates living amongst the foliage of oaks and other trees and shrubs. It was first discovered in the county at Forthampton Oaks (YC34; S083), with brown tree ant Lasius brunneus on the foliage of this large concentration of ancient oaks, 14.viii. 1 999. It has subsequently been found on the foliage of an ancient oak pollard at Newnham (VC34; S061), 5.viii.2004, and from pear foliage in an old orchard at Malswick, Newent (VC34; S072), 5.viii.2004. These are all sites with brown tree ant and its distribution pattern in the county closely fits the distribution of this ant.

The widespread southern flower bug Orius laticollis (Reut.) (Anthocoridae) was also discovered at Ashleworth Ham, 4.viii.2002, and is mostly associated with larger trees of poplar, willow and ash. It has also turned up at Debdene (VC33; SP12) in the Cotswolds, 17.vi.2000, and so appears not to be a floodplain speciality in the county.

The grass bug Miridius quadrivirgatus (Costa) (Miridae) was said to be mainly associated with wall barley where it grows in rough pastures near the sea (Southwood & Leston, 1959) but it has since then spread well inland and northwards

36

BR. J. ENT. NAT. HIST., 19: 2006

(B.S. Nau, pers.comm.). A single specimen was swept from the old grassland of Shuthonger Common (VC33; S083), 14.viii. 1999. It has subsequently been swept from ditch-side rough grassland at Sherborne Farm water meadows (VC33; SP1 1) in the Windrush Valley of the Cotswold Hills by John Widgery, 24.viii.2000.

Limestone grasslands

The third major geographical feature of Gloucestershire is the Cotswold Hills, with their large expanses of unimproved limestone pastures, large areas of enclosed ancient woodland, and a landscape of large old open-grown native trees. Although this area is probably the most favoured by recorders it has still generated the largest number of additional species, suggesting that the whole county remains under- recorded for Heteroptera.

The most significant find has probably been the seed bug Megalonotus antennatus (Schilling). It is not particularly associated with limestone grasslands (Kirby, 1992) but has only been found so far on one of the highest quality examples, Swifts Hill (VC33; SO80), a GWT reserve. One was found in a disused quarry there, another on the south-facing hillside, 19.vi.2003. A single specimen of another seed bug Peritrechus nubilus (Fallen) was found in Three Grove’s Wood GWT Reserve, Oakridge (VC33; SO90), 13.xi.2003, but this normally open country species is presumed to be an overwintering individual here from the neighbouring limestone pasture of Strawberry Banks GWT Reserve.

The stilt bug Berytinus crassipes (Herrich-Schaeffer) (Berytinidae) has a mainly eastern distribution in England, where it favours sparse but grassy wastes, especially old mineral workings, cinder railway embankments, feeding on mouse-ear chickweeds and other plants (Southwood & Leston, 1959). One was swept in an area of old quarry workings at Kilkenny Viewpoint (VC33; SP01), 8.vi.2002, and another was found on the old clinker bed of Chedworth Railway GWT Reserve (VC33; SP01), 23.vii.2003.

The Cotswold grasslands also include areas of unimproved neutral grassland and the knapweed associated plant bug Oncotylus viridiflavus (Goeze) (Miridae) was found at Box Farm Meadows (VC34; SO80), 23.vii.2003. Gloucestershire is close to the northern edge of its British range (Southwood & Leston, 1959).

The final open country species added to the county list from the Cotswolds is the lace bug Agramma laeta (Fallen) (Tingidae). It is thought to be associated with sedges, woodrushes and rushes, and occurs on saltmarshes, limestone grasslands, in damp woodland, and elsewhere. It is a typical inhabitant of old limestone grasslands and has been found widely on the old commons around Stroud (SO80): Minchinhampton Common, 3.vi.l985; Littleworth Common, 25.iii.1988; Selsley Common, lO.viii. 1 997 (all VC 34), and Swifts Hill, 18.vi.2003. Andy Foster (pers. comm.) has also taken it on Haresfield Beacon (VC33), 28.vii.1998. One further site has also been found on the edge of the Forest of Dean in the old limestone quarry Stenders Quarry (VC34; S061), 5.viii.2003.

Two further additions come from tree and woodland sites on the Cotswolds. A specimen of the nationally scarce plant bug Psallus albicinctus (Kirschbaum) was swept from grassland downwind of old boundary oaks near Old Hinchwick (VC33; SP12) on a very windy day, 17. vi. 2000. It has a south-eastern distribution in England, from Dorset to Northamptonshire, and is associated with mature oaks in open-structured woodland or wood-pasture trees, possibly specialising in high canopy (Kirby, 1992).

BR. J. ENT. NAT. HIST., 19: 2006

37

Another plant bug, Mecomma dispar (Boheman) is also a surprising addition to the list. Although a boreo-montane species, this bug has an odd distribution in Britain involving the north and west -as might be expected but also Surrey, East Anglia and a few Midland counties, but is rare in the South West (Southwood & Leston, 1959). Its habitat associations are also diverse, including rank vegetation, on marshes, on sand dunes, and also in short turf on sea cliffs. One was swept along a wide grassy ride in Siccaridge Wood (VC33; SO90), ll.vi.2003.

ACKNOWLEDGEM ENTS

I would like to record my thanks to Colin Twissell and other members of the Gloucestershire Invertebrate Group who have accompanied me on recording trips and often found the more interesting bugs for me. Access was usually arranged by Rosie Cliffe of the Gloucestershire Wildlife Trust, who also commissioned some reserve surveys during 2003, funded by the Heritage Lottery Fund. Thanks also to Bernard Nau and Pete Kirby for help with determinations; and to Andy Foster and John Widgery for passing on their records.

References

Alexander, K.N.A. 1995. The land and freshwater bugs (Hemiptera) of Gloucestershire: Part 1. Shield bugs to Stilt bugs (Heteroptera: Pentatomorpha). The Gloucestershire Naturalist 8: 37-53.

Alexander, K.N.A. 1996. The land and freshwater bugs (Hemiptera) of Gloucestershire: Part 2. Lace bugs to plantbugs (Heteroptera: Cimicomorpha). The Gloucestershire Naturalist 9: 31-62.

Kirby, P. 1992. A review of the scarce and threatened Hemiptera of Great Britain. Joint Nature Conservation Committee: UK Nature Conservation No. 2.

Southwood, T.R.E., & Leston, D. 1959. Land and Water Bugs of the British Isles. Warne, London.

SHORT COMMUNICATIONS

Schreckensteinia festaliella (Hiibner) (Lepidoptera: Schreckensteiniidae) on an unusual food-plant in the Outer Hebrides. - On 30.vii.2005 in the gorge-woodland near the mouth of Allt Volagir (O.S. Grid NF7929), on South Uist (VC1 10), many of the hazel trees ( Corylus avellana ) had marked “browning” of the leaves. Closer inspection revealed that this browning took the form of many small elongate areas of windowing, due to larval feeding removing portions of the upper epidermis parallel to the larger veins. Careful searching soon revealed the characteristic green larva of Schreckensteinia festaliella feeding exposed on the upper surface of the leaf. These larvae subsequently formed the typical spindle-shaped network cocoons of S. festaliella.

In my experience in Scotland, S. festaliella larvae normally feed on raspberry, Rubus idaeus, and less frequently on bramble, R. fructicosus agg. and stone bramble, R. saxatilis. On South Uist, the brambles under and near the infested hazel trees had signs of S. festaliella larval feeding, but to a far lesser extent than the hazel bushes. This report appears to be the first record of S. festaliella from the Outer Hebrides (YC1 10) and the first report of hazel as a food-plant for this species. - K. P. BLAND, National Museums of Scotland, The Granton Centre, 242 West Granton Road, Edinburgh, EH5 1JA.

38

BR. J. ENT. NAT. HIST., 19: 2006

Mortality of Orthoptera caused by mechanised mowing of grassland. - Orders such as Orthoptera are prey for bird and spider species (Joern, 1986; Belovsky & Slade, 1993; Oedekoven & Joern, 1998) and are known to be an important component of the diet of late season Cirl bunting Emberiza cirlus chicks (Evans et al., 1997). Therefore the decline of this species may be directly attributable to the loss of invertebrate diversity and abundance, particularly of Orthoptera, from farmland. Gardiner et al. (2002) and Gardiner & Hill (2003) found that Orthoptera were scarce in many agricultural grasslands that were heavily grazed or cut for silage, perhaps because of the short swards created by such management, which do not afford grasshoppers shelter from predation or inclement weather. However, the physical process of silage cutting is likely to have significant effects on grasshoppers that are in the sward when mowing occurs.

A study by Gardiner & Hill (2005) found that male and female Meadow Grasshopper Chorthippus parallelus Zetterstedt adults resided for the majority of time in the ground zone (sward height: <20 cm) and that this may lead to high mortality when grass cutting occurs. Grasshoppers are however very mobile and can jump large distances to escape from predators (Clarke, 1948; Richards & Waloff, 1954) and as a consequence may be able to escape damage or death during harvesting of silage.

Further to the study outlined in Gardiner & Hill (2005), laboratory examinations of cut hay (on 10 July 2004) and silage herbage (on 20 May and 10 July 2004; two-cut strategy determined by local climate and soil conditions) on the Writtle College Estate (site grid reference: TL664067) were undertaken to determine whether mechanised cutting caused mortality of Orthoptera. The hay and silage treatments were cut using a ride-on rotary mower (cutting height: 90 mm). Herbage from 80 randomly located 10 x 10 cm quadrats was collected from the hay and silage treatments (total area of herbage collected for each treatment: 0.8 m2) two hours after cutting. The cut herbage within the quadrat boundaries was collected and transported back to the laboratory in plastic bags. The contents of each sample were then emptied onto a white tray and searched for two minutes in an effort to locate dead and live Orthoptera.

Three dead adult C. parallelus individuals were found in the hay herbage collected on 10 July (Table 1). Mowing on 10 July also caused mortality of nymphs, two dead C. parallelus nymphs each were collected from the hay (both fourth instar) and silage (both second instar) treatments. A dead third instar nymph of the Lesser Marsh Grasshopper C. albomarginatus (De Geer) was found in the silage herbage on 10 July, but no dead or live individuals of any species were collected from the silage herbage after cutting on 20 May. In contrast to recording dead C. parallelus individuals, a total of five live adults (3 2 3) of this species were collected from the

Table 1 . Total number of dead and live grasshoppers in herbage of the hay and silage treatments mown on 10 July 2004

Species/life stage	Hay		Silage	Total
Dead	Live	Dead	Live	Dead Live
Chorthippus parallelus adult	3	2	2	3	5 5
C. parallelus nymph	2	0	2	1	4 1
C. albomarginatus nymph	0	0	1	1	1 1

BR. J. ENT. NAT. HIST., 19: 2006

39

cut herbage, of which three were collected from the silage treatment. One live nymph of both C. parallelus (second instar) and C. albomarginatus (second instar) were also observed in the cut silage herbage.

Although only a small scale investigation of cut hay and silage herbage, the study does show that Orthoptera of all developmental stages were killed by rotary cutting blades or contact with machinery during the process of hay and silage mowing. Dead individuals of two species were identified from the cut herbage in this experiment: C. albomarginatus and C. parallelus , both common grasshopper species on farmland at Writtle College (Gardiner et al. , 2002). The latter species is known to inhabit the ground zone (<20cm sward height) of hay meadows where it primarily exhibits resting and basking behaviour (Gardiner & Hill, 2005) making it vulnerable to disturbance and mortality from cutting blades which pass through the vegetation at approximately 10 cm height. The initial response to disturbance of species such as C. parallelus is to jump (Clarke, 1948); this means that individuals which become trapped under the mower are likely to jump into the rotary blades as an initial response to the disturbance caused by cutting. No assessment was made of the scale of mortality of grasshoppers on the swards, although it is likely that the mortality rate was fairly high as 10 dead individuals were identified from a relatively limited area of herbage (1.6 m2) with an average population density of 0.50 adult grasshopper ( Chorthippus spp.) individuals per m2 in the sward (Gardiner et al ., 2002).

Wagner (2004) concluded that mortality of Metrioptera bicolor Philippi due to cutting was high in German hay meadows (mortality rate: 42% of marked individuals). Other species such as Large Marsh Grasshopper Stethophyma grossum L. and Speckled Bush-cricket Leptophyes punctatissima Bose are also susceptible to injury or death by cutting blades (Oppermann & Krismann, 2001). These authors showed that injury rates of adult Orthoptera from rotary mowers were dependent on body size; larger individuals being more prone to injury. Therefore, large mature nymphs (fourth instar) and adults that are frequent in early July (Marshall & Haes, 1988) may be particularly prone to death or injury from hay and silage cutting that is undertaken at this time of year.

Perhaps as interesting as the confirmation of dead individuals from both hay and silage herbage, was the presence of live individuals. Live individuals of C. albomarginatus and C. parallelus were present in the cut herbage (Table 1), indicating that transfer of cuttings from one site to another (hay strewing) may transport species between sites. Wagner (2004) suggests that sexually active females of M. bicolor that survive cutting of hay can be transported to donor sites in the cut herbage. -Tim Gardiner, Centre for Environment & Rural Affairs (CERA), Writtle College, Chelmsford, Essex, UK, CM1 3RR & Julian Hill, Faculty of Land and Food Resources, University of Melbourne, Parkville, Victoria 3010, Australia.

References

Belovsky, G.E. & Slade, J.B. (1993). The role of vertebrate and invertebrate predators in a grasshopper community. Oikos 68: 193-201.

Clarke, E.J. (1948). Studies in the ecology of British grasshoppers. Transactions of the Royal Entomological Society of London 99: 173-222.

Evans, A.D., Smith, K.W., Buckingham, D.L. & Evans, J. (1997). Seasonal variation in breeding performance and nestling diet of Cirl Buntings Emberiza cirlus in England. Bird Study 44: 66-79.

Gardiner, T. & Hill, J. (2003). Are there any grasshoppers on farmland? Antenna 27: 115-116.

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BR. J. ENT. NAT. HIST., 19: 2006

Gardiner, T. & Hill, J. (2005). Behavioural observations of Chorthippus parallelus (Orthoptera: Acrididae) adults in managed grassland. British Journal of Entomology and Natural History 18: 1-8.

Gardiner, T., Pye, M., Field, R. & Hill, J. (2002). The influence of sward height and vegetation composition in determining the habitat preferences of three Chorthippus species (Orthoptera: Acrididae) in Chelmsford, Essex, UK. Journal of Orthoptera Research 11: 207-213.

Joern, A. (1986). Experimental study of avian predation on coexisting grasshopper populations (Orthoptera: Acrididae) in a sandhills grassland. Oikos 46: 243-249.

Marshall, J.A. & Haes, E.C.M. (1988). Grasshoppers and Allied Insects of Great Britain and Ireland. Harley Books, Colchester.

Oedekoven, M.A. & Joern, A. (1998). Stage-based mortality of grassland grasshoppers (Acrididae) from wandering spider (Lycosidae) predation. Acta Oecologica 19: 507-515.

Oppermann, R. & Krismann, A. (2001). Naturvertragliche Mahtechnik and Populationssicher- ung. Skripten des Bundesamtes fur Naturschutz 54: 1-76.

Richards, O.W. & Waloff, N. (1954). Studies on the biology and population dynamics of British grasshoppers. Anti-Locust Bulletin 17: 1-182.

Wagner, C. (2004). Passive dispersal of Metrioptera bicolor (Phillipi 1830) (Orthopteroidea: Ensifera: Tettigoniidae) by transfer of hay. Journal of Insect Conservation 8: 287-296.

Exochomus quadripustulatus (L.) (Coleoptera: Coccinellidae) as a host of Dinocampus coccinellae (Schrank) (Hymenoptera: Braconidae). - During an extended study of mortality of pine ladybirds Exochomus quadripustulatus (L.), mainly in Sheffield, the bulk of parasitoids that emerged from the ladybird pupae, were identified as the chalcid Aprostocetus neglectus (Domenichini) (Hymenoptera: Eulophidae). In early May, 2004 a pine ladybird imago was observed in a crack of bark on a sycamore tree in Millhouses Park (VC 63, SK3383) which was immobile for at least a week. It was removed on 18th May and seen to be attached to the tree by a cocoon. Kept in a fairly warm place, a wasp emerged on 24th May which appeared to be a small Dinocampus coccinellae (Schrank). This was confirmed by Dr Mark Shaw.

Majerus (1997, Br. J. Ent. Nat. Hist. 10: 15-24) detailed his observations of the parasitoid Dinocampus coccinellae predating various British coccinellid species. This parasitoid, which develops within the ladybird imago and pupates below it, was not observed on c. 3000 specimens of E. quadripustulatus nor any other members of the sub-family Chilocorinae. Majerus quotes the non-inclusion of any of the British chilocorines as hosts of the braconid in the list of Hodek (1973, Biology of Coccinellidae , Junk/ Academic Press). Neither do Klausnitzer & Klausnitzer (1977, Marienkafer, Westarp Wissenschaften, Magdeburg) include chilocorines as hosts. However, Hodek does include a record of D. coccinellae parasitizing Exochomus concavus Fursch in Transvaal, South Africa. This therefore appears to be the first record of E. quadripustulatus as prey and possibly the first palaearctic record from any chilocorine.

I thank Dr Shaw for his advice. The organisms and cocoon are deposited in the Royal Museum of Scotland. - Paul R. Mabbott, 49 Endowood Road, Millhouses, Sheffield, S7 2LY. mabbott@blueyonder.co.uk

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ON THE IDENTITY OF PSEUDEXECHIA PARALLELA (EDWARDS, 1925) (DIPTERA: MY CETO PHILID AE) AND DESCRIPTION OF A NEW RELATED SPECIES FROM GREAT BRITAIN

JOSTEIN KLERANDSEN & PETER J. CHANDLER*

Museum of Zoology, Lund University, Helgonavdgen 3, S-223 62 Lund, Sweden. E-mail: jostein.kjaerandsen@zool.lu.se.

*606B Berryfield Lane, Melksham, Wilts SN12 6EL, U.K. E-mail: Chandgnats@aol.com.

Abstract

Pseudexechia parallela (Edwards, 1925) is redescribed and illustrated. Based on study of types and new material Pseudexechia parallela (Edwards, 1925) is considered to be a senior synonym of P. hamulata (Lackschewitz, 1937) syn. n. Pseudexechia parallela is reported from Sweden, Kamtschatka, Russia and Minnesota, U.S.A. for the first time, thus expanding its known range from Europe to a wide Holarctic distribution. A new allied species, Pseudexechia monica sp. n., is described and illustrated based on males from Wales in Great Britain.

Introduction

The genus Pseudexechia Tuomikoski was established by Tuomikoski (1966) for a small group of closely related species segregated from Exechia Winnertz, characterised by the absence of discal bristles on the mesoscutum, ovate clypeus and distinctive features of the male terminalia such as the bud-like hypandrial lobe (Chandler, 1978). At present there are 22 known species associated in this genus from the Holarctic, the Oriental and the Afrotropical regions. However, several species await description and some that were described in other genera are yet to be formally transferred to Pseudexechia.

Pseudexechia parallela (Edwards, 1925) was described based on a single British female with a two-segmented cercus. Edwards (1925) considered it to be an easy task to associate it to the male based on key characters such as parallel branches of the cubital fork and a straight Rs vein. However, the association proved to be problematic as the wing venation of the holotype is apparently rather aberrant with respect to the narrow cubital fork. Consequently, Lackschewitz (1937) described a second very close species, P. hamulata (Lackschewitz, 1937), from Latvia without any notes on its relationship to P. parallela. Stackelberg (1948) subsequently associated and figured the male of P. parallela based on material from the Leningrad district of Russia. Burghele-Balacesco (1972) reported P. parallela from Romanian caves and was the first to notice that the narrow cubital fork was not a constant character. Chandler (1978) reviewed the known Holarctic species of Pseudexechia without adding any material or further description of P. parallela. Pseudexechia parallela was later incorrectly associated and figured by Krivosheina et al. (1986), whose figure represents another species of which the identity is yet to be confirmed. Zaitzev (2003) questioned the validity of P. hamulata as separate from P. parallela , referring to Chandler (1978). At present P. parallela is reported to be widespread in most of Europe while P. hamulata is known only to have a disjunct distribution in Estonia, Latvia and Denmark (Chandler, 2004).

No clear difference is apparent between the two species based on previously published illustrations. At most it seems that P. hamulata as figured by Lackschewitz (1937) has a sharper apicodorsal corner of the ventral branch of the gonostylus than

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has P. parallela as figured by Stackelberg (1948). This character, however, is liable to great variation in dorsoventral view depending on the exposed angle of the ventral branch, and only the internal face of the gonostylus will reveal the true shape of this thin plate (not figured by Stackelberg 1948).

New material of Pseudexechia from Great Britain has revealed two closely related species of which one is more widespread and represented by both sexes. We consider this to be the true P. parallela , thus confirming the identity of its male. When compared with the holotype of P. hamulata we find the two to be identical. Hence, we consider P. parallela to be a senior synonym of P. hamulata. Unfortunately we have not been able to re-examine the material that was studied by Stackelberg (1948), but based on his figures we find it very likely that this material was correctly assigned to P. parallela. Other British specimens represent a new species, so far known only from Wales. A review of all Nordic species of Pseudexechia will be presented elsewhere (Kjaerandsen in prep.).

Material and methods

The following abbreviations are used for institutions in which specimens are deposited. Other specimens are in the private collection of P. J. Chandler.

BMNH - Natural History Museum, London, England.

FMNH - Finnish Museum of Natural History, Helsinki, Finland.

MZLU - Museum of Zoology, Lund, Sweden.

ZMHB - Museum fur Naturkunde der Humboldt-Universitat, Berlin, Germany.

The general terminology follows Soli (1997), the measurements and ratios follow Kjaerandsen & Kurina (2004), and the terminology of the male gonostylus follows Kjaerandsen (in press). In cases where a range of measurement is given, the figure following the range is the mean.

The species

Pseudexechia parallela (Edwards, 1925)

(Fig. 1)

Exechia parallela Edwards, 1925: 596.

Pseudexechia hamulata (Lackschewitz, 1937), syn. n.

- Type material examined: Holotype (female) of P. parallela : ENGLAND (U.K.): Newmarket, Cambs, 23 Sep 1888 (coll. G. H. Verrall) - SPM-005283 (BMNH, pinned with terminalia in balsam preparation mounted on pin); holotype (male) of P. hamulata: LATVIA: Paplacken, Kurl, 7 Oct 1934 (coll. P. Lackschewitz) - SPM- 011949 (ZMHB, pinned with terminalia in balsam preparation mounted on pin).

- Other material examined: ENGLAND (U.K.): (all NCC Wetland Survey, coll. A. Foster and D. Procter): Cambridgeshire, Chippenham Fen, 10-24 Aug 1988 - 2 males; Norfolk, 13 Jun-1 1 Jul 1988 - 5 males, 2 females (MZLU, 1 male, 1 female on slide); Reedham, 27 Jun 1 1 Jul 1988 - 1 male, 1 1 Jul-12 Aug 1988 - 1 male, 12 Aug- 20 Sep 1988 - 2 males (MZLU, 1 male on slide), 26 Aug 1988 - 3 males; Brancaster,

1 -15 Jul 1988-9 males (MZLU, 1 male pinned); Stallode Wash, 29 Jun-8 Jul 1988 - 1 male; Catfield, 29 Sep-12 Oct 1988 - 1 male, 12 Oct 1988 1 male; Middle Harling,

26 Jun-1 7 Jul 1988 - 1 male; Wendling, 14-24 Sep 1988 - 1 male; Old Buckenham Fen, 28 Jun 12 Jul 1988 1 male; Thompson Common, 1988 1 male; Suffolk,

Walberswick, 14-29 Aug 1988 - 4 males; WALES (all NCC Peatland Survey, coll.

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43

Holmes, Boyce & Reed): Anglesey, Cors Erddreiniog, 27 Jul 1988 - 2 males; Cors Bodeilio, Molinia / Myrica bog and Phragmites bed, 28 Jun 1988 - 1 female, 26 Jul 1988 - 1 female, 5 Oct 1988 - 1 male; Rhos-y-Gad, calcareous flush fen, 27 Jul 1988 1 male, 1 female; West Glamorgan, Fairwood Common, J uncus flush, 4 Oct 1989 - 1 female; Pant-y-Sais, among Car ex acutiformis and in Molinia bog, 5 Oct 1989 - 2 males, 1 female; SWEDEN: SK, Hackeberga Nature Reserve, 29 Sep 1988 (coll. L. Huggert) - 3 males (MZLU, two on slide); ESTONIA: Torva, Helme Cave [old refuge cave in sandstone, about 50 m deep], 5801'00"N, 02553'00"E, 19 Jan 1996 (coll. O. Kurina) - 2 males (MZLU, on slides); RUSSIA: Kamtschatka, Bolscherjetsk, 1 Jul 1917 (coll. Y. Wuorentaus) - 1 male (FMNH, pinned with cleared terminalia in glycerine); USA: Minnesota, Cook co., Hovland N. J., Min. F. S„ 14 Jul 1968 (coll. E. F. Cook) - 1 male (MZLU, on slide).

Diagnostic characters: P. parallela forms together with P. monica sp. n. a group (also including an undescribed Oriental species) of small Pseudexechia species with dark brown mesoscutum bearing fused thoracic stripes, and with reduced size of the gonostylus relative to the gonocoxite (normally protruding less than a fifth gonocoxal length beyond its apex depending on orientation of gonostylus). The latter character also applies to P. aurivernica Chandler 1978 but that has distinct thoracic stripes. It can be separated from P. monica by unique characters in the male terminalia, including the ventral branch of gonostylus that is apicodorsally pointed, the hypandrial lobe being concave with a small, slightly diverging split apically, and the hypoproct forming a long, only slightly down-curved cylindrical process. Pseudexechia parallela is so far the only known species of Pseudexechia where the female has a two-segmented cercus (females of a second species not yet associated with males is known from Sweden, but this has distinct thoracic stripes so is not considered to be P. monica).

Description

Male ( n = 6 , except where otherwise stated). Total length 3. 7-4.8, 4.3 mm. Wing length 2.56-3.2, 2.82 mm, or 2.98-3.38, 3.21 x as long as profemur. Mesoscutum length 0.66-0.82, 0.75 mm, or 0.25-0.28, 0.26 x as long as wing.

Coloration (dry, pinned specimens, n = 42). Head brown, grey dusted. Antenna with scape, pedicel and first flagellomere yellowish brown, otherwise darker brown. Palpus brownish yellow. Thorax with mesoscutum mostly dark brown and grey dusted on disc without separate stripes, leaving only the side margins and humeral areas yellowish. Dark coloration continued onto disc of scutellum. Pleura yellowish brown. Legs yellow, including tibial spurs. Wing yellowish, with costa and radial veins slightly darker. Abdomen with tergites mostly dark brown dorsally, but tergites 1-5 with yellow lateral markings, more or less triangular on 2-5 and broadest on hind margins. Tergite 6 all brown. Sternites and genitalia brownish yellow.

Head. Round, width / length to frontal tubercle 1.4-1.48, 1.44. Antenna 1.34-1.66, 1.51 mm long. First flagellomere 1.67-2.04, 1.85 times as long as second flagellomere. Second flagellomere 1.29-1.94, 1.66 times as long as wide. Two ocelli present, touching compound eyes. Clypeus ovate, length/width 1.1-1.28, 1.18. Antepenulti- mate segment of maxillary palp 0.10-0.11, 0.10 mm long, palpomere ratios 1: 1.11- 1.44, 1.3: 1.83-2.62, 2.42.

Thorax. Proepisternum with one strong and one smaller bristle. Mesoscutum with some strong front-marginal, prealar and postalar bristles, otherwise covered with small dark, decumbent setae. Scutellum with one pair of strong bristles.

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Fig. 1. Terminalia of Pseudexechia parallela (Edwards, 1925). A. Male terminalia, ventral view; B. Internal face of gonostylus, enlarged; C. Male tergite 9, cerci and hypoproct, dorsal view; D. Female terminalia, lateral view.

Wings. Costa and R5 with setae both dorsally and ventrally, stem of R and Rj with dorsal setae only, except a few (0-15, 7) ventral setae apically on Rj. Rs, ta, tb, M and Cu without setae. Wing length to length of Rj 2. 2-2.4, 2.32. R5 nearly straight, wing length to length of R5 1.73-1.77, 1 .75. Length of ta to length of M-petiole 0.89-

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1.11, 1. Cu-fork short, fork length ratio (A/B) 0.76-0.8, 0.78. Fork width ratios (C/D and E/F) 1.14—1.28, 1.21 and 0.8-1.2, 0.98. M-ratios 0.58-0.63, 0.6 and 0.64-0.7, 0.66. CuA-ratios 1 .35-1 .53, 1 .45 and 1.81-2.11, 1 .99. Cubital fork width (F) to length of CuAj 0.33-0.41, 0.36. CuP vague, ratio of length to length of wing 0.41-0.49, 0.45. A! distinct, ratio of length to length of wing 0.34—0.38, 0.36.

Legs. Leg ratios given for fore, mid and hind leg: LR 1.11-1.17, 1.14: 0.81-0.86, 0.84: 0.63-0.68, 0.65; SV 1.61-1.69, 1.65: 2.04-2.15, 2.08: 2.52-2.68, 2.61; BV 1.51- 1.65, 1.57: 1.87-2.05, 1.94 (n = 5): 2.76-3.1,2.98; TR 1.53-1.67, 1.58: 1.58-1.74, 1.67: 1.85-2.07, 1.96.

Terminalia. Tergite 9 (Lig. 1C) medially divided into semicircular plates, with setae of variable size. Cercus long, slender, 6-7, 6.4 as long as wide in dorsal view. A second small ovate, setose lobe present medial to cercus (this lobe of questionable origin but possibly resulting by separation from a bilobed cercus as found in other related genera). Gonocoxite (Fig. 1A) open dorsally and moderately incised ventrally, without medial suture. Hypandrial lobe concave, apically with a small, slightly diverging split. Hypoproct well developed, apically prominent, forming a long, only slightly down-curved cylindrical process, reaching beyond apex of cercus, ventrally forming rectangular recess, apparently fused with small conical aedeagal apparatus. Gonocoxal apodeme long, moderately sclerotised, apically broadened with two condyles. Accessory copulatory appendages hyaline and difficult to interpret. Gonostylus (Fig. IB) with five branches. Dorsal branch with strong setae on lateral face, apically truncated to whitish, smooth cushion, devoid of setae. Dorsointernal branch wide, nearly symmetrically fan shaped, with 23-24 lamellae. Medial branch reduced or absent. Ventral branch forming thin plate, apically widened, subcircular, apicodorsally prolonged into acute angled pointed corner; basally with one very long and one shorter seta, apicolateral face with 2^4 strong, fan- tipped setae and group of small regular setae; apicointernal face with some small setae. Internal branch forms a large, subrectangular pouch, partly striated, ventroapically with a few strong setae. Anterior branch present as large hyaline cushion, devoid of setae.

Female ( n = 1). Total length 4.4 mm. Wing length 2.84 mm, or 3.23 x as long as profemur. Mesoscutum length 0.8 mm, or 0.28 x as long as wing.

Coloration (n = 5). As for male except more extensive yellow markings on abdomen, yellow lateral triangles also on tergite 6, tergite 7 and ovipositor entirely yellow.

Head. Round, width / length to frontal tubercle 1.45. Antenna 1.4 mm long. First flagellomere 1.67 times as long as second flagellomere. Second flagellomere 1.67 times as long as wide. Two ocelli present, touching compound eyes. Clypeus ovate, length/width 1.14. Antepenultimate segment of maxillary palp 0.12 mm long, palpomere ratios 1: 1.18: 2.1.

Thorax. As for male.

Wings. Setation as for male, Rj with 8 setae apically. Wing length to length of Rj 2.29. R5 nearly straight, wing length to length of R5 1.67. Length of ta to length of M-petiole 1.11. Fork length ratio (A/B) 0.74. Fork width ratios (C/D and E/F) 1.28 and 1. M-ratios 0.58 and 0.63. CuA-ratios 1.56 and 2.19. Cubital fork width (F) to length of CuAj 0.38. CuP vague, length to length of wing 0.48. A! length to length of wing 0.35.

Legs. Leg ratios given for fore, mid and hind leg: LR 1.14: 0.87: 0.67; SV 1.64: 2.03: 2.56; BV 1.56: 1.99: 2.96; TR 1.61: 1.79: 2.03.

Terminalia (Fig. ID). Tergite 7 with straight margin apicodorsally, apicolateral margin slightly produced, evenly convex. Cercus two-segmented, scattered with small setae; apical segment ovate, twice as long as wide in lateral view. Gonapophysis 8

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broad, with slightly truncated apex, with gradually stronger setae towards apex. Postgenital plate stout, bluntly tapered, apically sclerotized and with numerous small setae.

Variation. The narrow cubital fork in the holotype, described as having parallel branches, is considered to be aberrant. The measured specimens all have distinctly divergent branches, but the cubital fork width (F) to length of CuA, ratio is rather variable [0.33-0.41, 0.36 (n = 7)], i.e. up to 24% wider in the specimen with the widest cubital fork compared to the specimen with the narrowest cubital fork. Some variation in the apicodorsal prolongation of the ventral branch of the male gonostylus is also notable, but considered to be within the species limits.

Distribution. Recorded from Sweden, Kamtschatka, Russia and Minnesota, U.S.A. for the first time, thus expanding its known range from most of Europe (see Chandler, 2004) to a wide Holarctic distribution. The British sites are a wide range of wetland habitats and the new material was obtained by water trapping in two major wetland surveys carried out by the former Nature Conservancy Council. It has not been found during other extensive collecting of Mycetophilidae in the British Isles in recent years but a single male was recorded from County Kerry in Ireland (Chandler et al., 2000; Falk & Chandler, 2005).

Pseudexechia monica sp. n.

(Fig- 2)

- Type material: Holotype male: GREAT BRITAIN (U.K.): WALES, Anglesey, Cors Erddreiniog, 27 July 1988, NCC Peatland Survey, water trap 9, rank Schoenus flushes, coll. Holmes, Boyce & Reed (deposited in National Museums of Scotland, Edinburgh).

Paratypes: GREAT BRITAIN (U.K.): WALES, same data as holotype 1 male; Anglesey, Cors Erddreining, NCC Peatland Survey (water trap 11), 27 July 1988 (coll. Holmes, Boyce & Reed) - 1 male (MZLU, on slide); Anglesey, Cors Bodeilio, NCC Peatland Survey (water trap 1 - Phragmites bed), 26 July 1988 (coll. Holmes, Boyce & Reed) - 3 males (MZLU, 1 male on slide); Anglesey, Cors Bodeilio, Mar 1990 (coll. A, Godfrey) - 1 male.

Etymology: The name is an adjective referring to the origin of the type material from the Isle of Anglesey, known in Latin as Mona.

Diagnostic characters: P. monica forms together with P. parallela a group of small Pseudexechia species with dark brown mesoscutum bearing fused thoracic stripes and with reduced size of the gonostylus relative to the gonocoxite. It can be separated from P. parallela by unique characters in the male terminalia, including the ventral branch of the gonostylus being angled and club shaped, the hypandrial lobe being large subrectangular, elaborate and with a large diverging split apically, and the hypoproct forming a shorter, strongly downcurved cylindrical process. It shares the shape of the ventral branch of the gonostylus with P. aurivernica but can easily be separated from this larger species by fused thoracic stripes and other details of the male terminalia such as shape of the hypandrial lobe and the widened apex of the dorsal branch of the gonostylus.

Description

Male (n = 2, except where otherwise stated). Total length 3. 9-3. 9 mm. Wing length 2.58-2.6 mm, or 2.89-3 x as long as profemur. Mesoscutum length 0.7-0.74 mm, or 0.27-0.28 x as long as wing.

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Fig. 2. Male terminalia of Pseudexechia monica sp. n. A. Terminalia, ventral view; B. Internal face of gonostylus, enlarged; C. Tergite 9, cerci and hypoproct, dorsal view; D. Hypandrial lobe, ventral view, enlarged.

Coloration (< n = 7). Head brown. Antenna with scape, pedicel and first flagellomere yellowish brown, otherwise darker brown. Palpus brownish yellow. Thorax with mesoscutum mostly dark brown with grey dusting, with only the side margins and humeral areas yellowish. Dark coloration continued onto disc of scutellum. Pleura yellowish brown. Legs yellow, including tibial spurs. Wing yellowish, with costa and radial veins slightly darker. Abdomen with tergites mostly brown, but tergites 1-5 usually with yellow lateral markings, more or less triangular on 2-5 and broadest on hind margins but usually more restricted in extent than in P. parallela. Tergite 6 all brown. Sternites and genitalia brownish yellow.

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- Head. Round, width/length to frontal tubercle 1.31-1.36. Antenna 1.44- 1.62 mm long. First flagellomere 1.87-2.03 times as long as second flagellomere. Second flagellomere 1.72-1.88 times as long as wide. Two ocelli present, touching compound eyes. Clypeus ovate, length/width 1.25-1.28. Antepenultimate segment 0.1-0.11 mm long, palpomere ratios 1: 1.32- 1.44: 2.16-2.34.

Thorax. Proepisternum with one strong and one smaller bristle. Mesoscutum with some strong front-marginal, prealar and postalar bristles, otherwise covered with small dark, decumbent setae. Scutellum with one pair of strong bristles.

Wings. Costa and R5 with setae both dorsally and ventrally, stem of R and R} with dorsal seta only, except a few (2-10) ventral setae apically on R^ Crossvein ta with one seta ventrally. Rs, tb, M and Cu without setae. Wing length to length of Rj 2.26-2.32. R5 nearly straight, wing length to length of R5 1.7-1.71. Length of ta to length of M-petiole 1.29-1.36. Cu-fork short, fork length ratio (A/B) 0.69-0.72. Fork width ratios (C/D and E/F) 1.15-1.22 and 1.08-1.19. M-ratios 0.54-0.56 and 0.59- 0.61. CuA-ratios 1.63-1.69 and 2.2-2.36. Cubital fork width (F) to length of CuAt 0.33-0.35. CuP vague, ratio of length to length of wing 0.48-0.5. Aj distinct, ratio of length to length of wing 0.33.

- Legs. Leg ratios given for fore, mid and hind leg: LR 1.15-1.16: 0.87-0.91: 0.65- 0.66; SV 1.6-1.65: 1.97-2.02: 2.58-2.63; BV 1.53: 1.87-1.88: 2.67-2.76; TR 1.54-1.61: 1.66-1.68: 1.78-1.84.

- Terminalia. Tergite 9 (Fig. 2C) medially divided into semicircular plates, with setae of variable size. Cercus long, slender, 4.67-6.18 as long as wide in dorsal view. A second small ovate, setose lobe present medial to cercus (of uncertain origin as indicated under P. parallela). Gonocoxite (Fig. 2A) open dorsally and deeply incised ventrally, with medial suture. Hypandrial lobe (Fig. 2D) large, subrectangular, elaborately folded, apically with a large diverging split. Hypoproct well developed, apically less prominent forming distinctly down-curved cylindrical process, not reaching beyond apex of cercus, ventrally forming rectangular recess, apparently fused with small conical aedeagal apparatus. Gonocoxal apodeme long, moderately sclerotized, apically broadened with two condyles. Accessory copulatory appendages hyaline and difficult to interpret. Gonostylus (Fig. 2B) with five branches. Dorsal branch with strong setae on lateral face, apically truncated and strongly widened to whitish, smooth cushion, devoid of setae. Dorsointernal branch narrowly fan shaped, with 17 large lamellae. Medial branch reduced or absent. Ventral branch forming thin plate, apically angled club shaped; basally with one very long and one shorter seta, apicolateral face with 3 regular, strong setae and group of small setae; apicointernal face with two strong and a few small setae. Internal branch forms a large, subrectangular pouch, partly striated, ventroapically with a few strong setae. Anterior branch present as large hyaline cushion, devoid of setae, apical rim sclerotized, with three tapered projections.

Female unknown.

- Distribution. Known only from Wales in Great Britain.

Acknowledgements

The study was financially supported by the Swedish Taxonomy Initiative (J. Kjaerandsen, see Miller, 2005). We are indebted to D. Notton (BMNH), P. Sihvonen (FMNH), R. Danielsson (MZLU), and W. Schacht (ZMHB) for the opportunity to work with the collections and for the loan of material.

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References

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Chandler, P.J. 1978. Notes on the Holarctic Species of Pseudexechia Tuomikoski (Diptera: Mycetophilidae), with the Description of a New British Species. Entomologist’s Record & Journal of Variation 90: 44—5 1 .

Chandler, P.J. 2004. Fauna Europaea: Mycetophilidae. In: de Jong, H. (Ed.) Fauna Europaea: Diptera, Nematocera. Fauna Europaea version 1.1. Available from: http://www.faunaeur. org (February 2005).

Chandler, P.J., O’Connor, J.P. & Nash, R. 2000. Diptera (Mycetophilidae, Empididae, Micropezidae, Drosophilidae, Sarcophagidae, Muscidae, Calliphoridae and Tachinidae) new to Ireland. Dipterists Digest (Second Series) 7: 15-18.

Edwards, F.W. 1925. British Fungus-Gnats (Diptera, Mycetophilidae). With a revised Generic Classification of the Family. Transactions of the Entomological Society of London 73: 505- 670.

Falk, S.J. & Chandler, P.J. 2005. A review of the scarce and threatened flies of Great Britain. Part 2: Nematocera and Aschiza not dealt with by Falk (1991). 187 pp. Joint Nature Conservation Committee, Peterborough.

Kjaerandsen, J. (In press): Review of fungus gnats of the genus Tarnania Tuomikoski, with a phylogeny of the Rymosia s.l. genus group (Diptera: Mycetophilidae). Insect Systematics and Evolution.

Kjaerandsen, J. & Kurina, O. 2004. A new species of Cordyla Meigen from Norway (Diptera: Mycetophilidae). Norwegian Journal of Entomology 51: 137-143.

Krivosheina, N.P., Zaitzev, A. I. & Yakovlev, E.B. 1986. [Insects as decomposers of fungi in the forest of the European part of USSR]. 309 pp., Moscow, [in Russian]

Lackschewitz, P. 1937. Die Fungivoriden des Ostbaltischen Gebietes. Arbeiten des Natur- forscher-Vereins zu Riga, Neue Folge 21: 1-47.

Miller, G. 2005. Linnaeus’s Legacy Carries On. Science 307: 1038-1039.

Soli, G.E.E. 1997. The adult morphology of Mycetophilidae (s.str.), with a tentative phylogeny of the family (Diptera, Sciaroidea). Entomologica Scandinavica, Supplement 50: 5-55.

Stackelberg, A. A. 1948. New and poorly known species of Fungivoridae (Diptera) from Leningrad District. Entomologicheskoe obozrenie 30: 94-102. [in Russian]

Tuomikoski, R. 1966. Generic taxonomy of the Exechiini (Dipt., Mycetophilidae). Annales entomologici Fennici 32: 159-194.

Zaitzev, A. I. 2003. Fungus gnats (Diptera, Sciaroidea) of the fauna of Russia and adjacent regions. Part II. An International Journal of Dipterological Research 14: 77-386.

ANNOUNCEMENT

Harley Books are delighted to announce the sale of their series, The Moths and Butterflies of Great Britain and Ireland , to Apollo Books Aps, Kirkeby Sand T9, DK-5771 Stenstrup, Denmark; email: apollobooks@vip.cybercity.dk; website: www.apollobooks.com.

With three of the ten volumes yet to be published - Yols 5, 6 and 8 Apollo Books, who are leading publishers of multi-volume works on European and world entomology, plan to complete the series. Dr K.P. Bland, Pelham-Clinton Curatorial Fellow at the National Museums of Scotland, Edinburgh, an associate editor and a contributing author to two volumes in the series, has been appointed editor.

All standing orders are being passed to Apollo Books and henceforward all matters relating to sales of this work, including orders for published volumes, should be referred to them. Apollo Books will continue to use Harley Books’ distributors and therefore British orders will be despatched from the UK.

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BR. J. ENT. NAT. HIST., 19: 2006

RESULTS OF THE

2005 BENHS MEMBERS’ QUESTIONNAIRE

Mark G. Telfer

10, Northall Road, Eaton Bray, Dunstable, Bedfordshire, LU6 2DQ

In order to plan the development of the Society, the Council and Officers of the British Entomological & Natural History Society wanted to gain a clearer understanding of Members’ views about various aspects of the Society’s current activities. A questionnaire was initiated by Mike Wilson. The questionnaire was developed through several further drafts during 2005 by Mark Telfer in consultation with Council members and Officers. The final version of the questionnaire was posted to all individual members in September, including members living outside Britain. Members attending the Annual Exhibition and Dinner on 12 November were given another opportunity to fill out a questionnaire if they had not already done so.

A deadline of 30 November was set for return of questionnaires. In practice, all questionnaires received before Christmas were included in the analysis, and only a few were returned later than that.

The questionnaire responses were numerically coded where necessary and entered into a custom-built relational database.

Most of the 24 questions about the Society were of the multiple-choice type. Five questions (questions 3, 7, 13, 18 and 23) asked respondents to order items by interest, preference or importance (the most interesting/preferred/important item getting 1, the next getting 2, and so on until indifferent about the remaining options). Many respondents used ties in their responses. If more than one option was allocated the same level of preference, they were all scored the same, e.g. all Tst equal’ were scored as ‘1’. If preferences were scored for example as follows: 1 = , 1 = , 1 = , 2, 3 = , 3 = , 4, 5, they were recoded on data entry as follows: 1, 1, 1, 4, 5, 5, 7, 8.

The questionnaire also asked respondents to write down the county in which they live, and concluded with a blank space in which respondents were invited to add any further comments they wished to make about the Society. Finally, respondents were given the option of putting their name on the questionnaire.

Results

334 completed questionnaires were returned in time to be included in this analysis. This is a return rate of 39% of the 857 questionnaires that were mailed out. This is felt to be an excellent rate of return.

The remainder of this Results section incorporates the subheadings and questions of the original questionnaire, followed by summaries of the responses.

The British Journal of Entomology and Natural History.

1. “Most of the contents of the Journal interest me”.

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disagree

Most members tended to agree that the contents of the Journal are interesting, though this still leaves some room for improvement.

Agree Tend to Neither Tend to Disagree strongly agree agree nor disagree strongly

disagree

Opinions on this issue were quite varied with most respondents plumping for the middle option, neither agreeing nor disagreeing. There was a slightly greater tendency to agree with the proposal than to disagree but on balance there seems to be no strong mandate for the Society to spend more on colour plates in the Journal.

3. Please number the following Journal sections in order of interest to you (the most interesting item scores 1, the next most interesting scores 2, and so on until you are indifferent about the remaining options):

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BR. J. ENT. NAT. HIST., 19: 2006

Table 1. Summary of responses to question 3, listed in decreasing order of interest. For each of the eight options, the average (mean) score is given, along with a measure of the amount of variation amongst the scores (standard deviation), and the number of scores (N).

	Average	Amount of variation	Number of scores
Articles	1.6	1.2	320
Short communications	2.5	1.2	312
Field Meeting reports	3.5	1.6	288
Exhibition reports	4.1	2.1	259
Book reviews	4.3	1.7	285
Indoor Meeting reports	5.1	1.6	229
Announcements	6.0	1.7	223
Officers’ reports	6.5	1.6	202

On average, Articles were judged the most interesting, and Officers’ reports the least interesting. Opinions on the Exhibition reports were more varied than on any other Journal section (note that the standard deviation of 2.1 is higher than for all other options), with some readers rating this very high and others very low.

Book publishing

4. Did you know that members are entitled to a third off Society publications?

Yes Not sure No

There was poor awareness of the discount available to members purchasing the Society’s books.

Indoor Meetings in London

The Society holds Indoor Meetings in London six times a year on Tuesday evenings at 18.00h.

5. How would you feel about a proposal to hold Indoor Meetings elsewhere in Britain?

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Approve Tend to Neither Tend to Disapprove strongly approve approve nor disapprove strongly

disapprove

There was quite a spread of opinion on this question. Attendances at the Indoor Meetings in South Kensington in recent years have been consistently disappointing. The bulk of the membership support the proposal to hold meetings elsewhere in Britain and Council should give this further consideration.

6. How would you rate the Indoor Meetings programmes?

Not Not very Quite Very Extremely

interesting at interesting interesting interesting interesting all

This question was intended to gauge whether the poor attendance at Indoor Meetings was caused by a lack of interest in the talks on offer, as opposed to the time, expense and inconvenience of attending on a Tuesday evening in London. It may not have done this very successfully and might have been better worded as: “How would you rate the talks on the Indoor Meetings programme (irrespective of whether or not you can attend)?”. Taken at face value, the responses appear to

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indicate that the talks on the programme are only ‘quite interesting’ to most members, leaving substantial room for improvement.

7. Please number the following types of talk in order of interest to you (the most interesting item scores 1 , the next most interesting item scores 2, and so on until you are indifferent about the remaining options):

Table 2. Summary of responses to question 7, listed in decreasing order of interest. For each of the nine options, the average (mean) score is given, along with a measure of the amount of variation amongst the scores (standard deviation), and the number of scores (N).

	Average	Amount of variation	Number of scores
British insects	2.1	1.5	278
Ecological studies	3.1	1.9	260
Conservation	3.3	1.9	271
Recording scheme activities	3.9	1.9	247
Taxonomic studies	4.3	2.4	211
History of natural history	5.2	2.2	211
Foreign insects	5.3	2.2	188
Natural history other than insects	5.7	2.4	190
Foreign travel	6.6	2.2	157

British insects, Ecological studies and Conservation were the most interesting subject areas on average. The least interesting subjects on average were Foreign travel, Natural history other than insects and Foreign insects.

Field Meetings across the UK

8. On average, how often do you attend the Society’s Field Meetings?

Never Less than About one a 2-5 per year 6 or more once a year year each year

Most respondents had attended at least one of the Society’s Field Meetings but were typically attending less than one a year. The Council should consider how to

BR. J. ENT. NAT. HIST., 19: 2006

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improve the levels of participation in Field Meetings, including by expanding the number of Field Meetings still further, by inviting new field meeting leaders to come forward, and by making the process of getting a field meeting on the programme clearer.

9. “The Field Meetings do not usually interest me”

Agree Tend to Neither Tend to Disagree strongly agree agree nor disagree strongly

disagree

A positive response regarding the interest of the Field Meetings, though probably leaving some room for improvement.

10. “The Field Meetings cover a good geographical spread”

Agree Tend to Neither Tend to Disagree strongly agree agree nor disagree strongly

disagree

A positive response regarding the geographical spread of the Field Meetings. This was an encouraging result given the geographical spread of the Society’s members

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BR. J. ENT. NAT. HIST., 19: 2006

and their differing willingness and ability to travel to Field Meetings. Further improvements could be made by making it easier for anyone who feels their area has been neglected in recent Field Meetings programmes to step forward as a leader.

11. “The Field Meetings cover the taxonomic groups I am interested in”

Agree Tend to Neither Tend to Disagree strongly agree agree nor disagree strongly

disagree

A positive response regarding the taxonomic coverage of the Field Meetings. There is probably room for improvement by encouraging those who feel their taxonomic interests are under-represented to come forward and lead Field Meetings.

12. “The Field Meetings are led by knowledgeable, experienced and helpful leaders”

Agree Tend to Neither Tend to Disagree strongly agree agree nor disagree strongly

disagree

A clear and highly positive response which is a credit to all those who have led Field Meetings for the Society in recent years.

BR. J. ENT. NAT. HIST., 19: 2006

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13. Please number the following types of Field Meetings in your order of preference (the most preferred scores 1, the next most preferred scores 2, and so on until you are indifferent about the remaining options):

Table 3. Summary of responses to question 13, listed in decreasing order of preference. For each of the six options, the average (mean) score is given, along with a measure of the amount of variation amongst the scores (standard deviation), and the number of scores (N).

	Average	Amount of variation	Number of scores
Unknown sites	2.3	1.4	249
Threatened sites	2.6	1.3	237
Search for particular species	3.0	1.6	234
Helping recording schemes	3.1	1.6	237
Known sites	3.3	1.7	215
Making management recommendations	4.7	1.6	193

The average preference scores were fairly closely bunched, with the exception of ‘Making management recommendations’ which was clearly the least favoured option. Respondents clearly favoured exploring unknown sites over known sites but also appreciated the role of Field Meetings in surveying threatened sites, searching for particular species and helping recording schemes.

The website: www.benhs.ovg.uk

14. Were you aware that the Society has a website?

Yes Not sure No

Most respondents were aware of the Society’s website. It seems likely that a substantial proportion of those who were unaware of the website or unsure of its existence are not internet users.

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15. On average, how often do you use the website?

It seems that the Society’s website is relatively little used, with only 14 respondents having used the website at least once a month.

16. If you have used the website, how would you rate it?

Only 137 respondents (41%) felt able to offer an opinion on the website; presumably most others had not visited the website. Most of those who did express an opinion felt the website was satisfactory.

There is considerable scope to improve the website and to encourage greater use of it and greater awareness of it.

BR. J. ENT. NAT. HIST., 19: 2006

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The Annual Exhibition and Dinner

17. Which of the following best describes your attendance at the Annual Exhibition in London?

years year

The number of respondents who attend the Annual Exhibition more often than not, is slightly outweighed by the number who never come or miss the exhibition more often than not. This represents something of a division in the Society. On the one hand there are many for whom the Exhibition is an important event in the Society’s calendar. Of the remainder, there may be some who are not interested in the Exhibition, though many wrote comments on their questionnaires to explain that they would like to attend but find the travel too expensive, awkward or time-consuming.

18. The council is considering changes to the Annual Exhibition venue which could give the Society much better value for money. Please number the following in order of importance to you (the most important thing scores 1 , the next most important scores 2, and so on until you are indifferent about the remaining options):

Table 4. Summary of responses to question 18, listed in decreasing order of importance. For each of the five options, the average (mean) score is given, along with a measure of the amount of variation amongst the scores (standard deviation), and the number of scores (N).

	Average	Amount of variation	Number of scores
Good public transport access	1.7	1.0	245
A more central location within Britain	2.1	1.3	200
Cheaper parking	2.4	1.2	193
Better lighting of the exhibition venue	3.2	1.1	150
A venue with a bar	3.9	1.2	151

Good public transport access was clearly the most important aspect of the Exhibition venue and the current venue at Imperial College, South Kensington does have very good public transport links. However, if an alternative venue could be found at a more central location within Britain with cheaper parking and equally good public transport links, this could prove more popular.

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Many respondents were indifferent about the lighting of the venue and the presence of a bar (note the much smaller number of respondents who scored these options).

Council should investigate alternative venues but members can be assured that Council recognises that many members are more than happy with the current arrangements: any changes will only be made after due consideration.

19. How would you feel about selected book dealers and equipment sellers being invited to the Exhibition?

Approve Tend to Neither Tend to Disapprove strongly approve approve nor disapprove strongly

disapprove

This proposal met with strong approval, providing one of the clearest results of the questionnaire. Council thus have a strong mandate to move forward with this proposal.

The Society’s Collections and Library are housed at Dinton Pastures, Reading

20. Were you aware of the facilities at Dinton Pastures?

Yes

Not sure

No

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There was excellent awareness of the Society’s facilities at Dinton Pastures.

21. On average, how often do you attend the Society’s Open Days at Dinton Pastures?

22. On average, how often do you attend the Society’s Workshops at Dinton Pastures?

Despite the excellent awareness of the Society’s facilities at Dinton Pastures, a relatively small proportion of members make use of those facilities, whether for the

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Open Days or the Workshops. Although members do travel from as far afield as Scotland and Devon to attend Dinton Pastures, it would seem that many members are put off by the journey. It may be worth noting here that Dinton Pastures is a simple 2 km walk from Winnersh station as it may not be widely appreciated just how easy it is to get to by public transport.

The Society as a whole

23. Please score the Society’s activities in order of importance to you. Give T’ to the most important, ‘2’ to the second most important, and so on to the end or until you are indifferent about the remainder.

Table 5. Summary of responses to question 23, listed in decreasing order of importance. For each of the nine options, the average (mean) score is given, along with a measure of the amount of variation amongst the scores (standard deviation), and the number of scores (N).

	Average	Amount of variation	Number of scores
Journal	2.0	1.4	318
Field Meetings	3.4	1.7	247
Books	3.7	1.9	246
Annual Exhibition and Dinner	3.8	2.7	216
Dinton Pastures (Collections and Library)	3.8	2.3	224
Workshops	4.2	1.9	219
Indoor Meetings	4.8	1.8	192
Public Liability Insurance of £5,000,000	6.5	2.9	147
Website	6.6	2.2	146

The nine options in this question represent the full range of the Society’s activities, of which the Journal was clearly felt to be the most important. Field Meetings were ranked as the second most important of the Society’s activities despite the fact that 39% of respondents had never attended a Field Meeting (question 8). The Society’s Books were ranked third, and like the Journal, this is an aspect of the Society’s activities that is accessible to all, irrespective of geographical location. By contrast, the Annual Exhibition and Dinner ranks fourth on average but the large variation in scores reveals the division in attitudes to the Exhibition noted under question 17. The largest variation was in scores for Public Liability Insurance. Various marginal comments revealed that many respondents were simply unaware of this insurance and thus could not offer an opinion on its importance: this needs to be better publicised.

24. “The subscription for 2006 of £19 a year provides good value for money”. Many Life Members felt unable to answer this question!

BR. J. ENT. NAT. HIST., 19: 2006

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disagree

There was a strong positive response that the Society does provide good value for money -one of the strongest positive responses from the questionnaire. This was a particularly welcome result considering that the questionnaire was issued shortly after the announcement of a rise in subscription rates.

Concluding Remarks

Of the 334 returned questionnaires, 192 (57%) included comments. These comments have been very helpful in interpreting the numerical results of the questionnaire presented here.

Many respondents took the opportunity to express their appreciation of the Society and the work done by its Council and Officers, such as “A pleasure to belong to such a well run Society”. These comments have been gratefully received.

Many respondents expressed their views on the logistical difficulties of participating in all the Society’s activities, such as the following from a member in Devon: “I think the BENHS an excellent society. The only downside is not symptomatic of the Society but rather of modern life (or just my lifestyle). The workshops, meetings and other interactive functions of the society are great but it is a case of whether one can make it to them. Keep up the good work”.

Many of the comments provided valuable detail on some of the problems encountered by members, often with constructive suggestions, all of which will help Council to decide on its responses to these results.

Acknowledgements

Thanks to all those who took the time to fill out and return the questionnaire. The excellent response rate is a testament to just how much members care about our Society. Particular thanks are also due to David Young for printing and mailing out the questionnaire. John Badmin and Mike Wilson commented on an earlier draft.

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BR. J. ENT. NAT. HIST., 19: 2006

ANNOUNCEMENT

Lepidoptera recording in northwest Wales (Merionethshire, Caernarvonshire and Anglesey). - In 2001 a grant from the BENHS Research Fund helped to establish a moth recording project in northwest Wales (VCs 48, 49, and 52). The underlying aim of the project is to make the best use of the efforts of a small number of dedicated resident recorders, by stimulating recording, improving data capture, and providing a report which acts as a reference to the range of species currently being recorded. So far we have seen production of 3 editions of the annual Lepidoptera report, improving co-ordination between Vice-county recorders, processing of part of the important Treborth dataset, and the ongoing development of Andrew Graham’s excellent database program for the area. Recent developments include a number of changes of ‘personnel’ in the Vice-county Recorder roles as listed below:

VC48 Merioneth

Moths & butterflies: Andrew Graham, Trawscoed, Llanuwchllyn, Bala, Gwynedd

LL23 7TD; e-mail: angrhm@globalnet.co.uk

VC49 Caernarfon

Moths: Julian Thompson, Pensychnant, Sychnant Pass, Conwy, Gwynedd, LL32 8BJ;

e-mail : julian @pensychnant . f snet .co.uk

Butterflies: David Thorpe, 3 Brynteg, Clwt y Bont, Gwynedd, LL55 3DT;

e-mail: david.thorpe@environment-agency.wales.gov.uk

VC52 Anglesey

Moths: John Harold, Hen Ardd, Carreg y Gath, Rhiwlas, Bangor, Gwynedd LL57

4HD; e-mail: jhmoths@yahoo.co.uk

Butterflies: David Thorpe, 3 Brynteg, Clwt y Bont, Gwynedd, LL55 3DT;

e-mail: david.thorpe@environment-agency.wales.gov.uk

As the new Moth Recorder for VC52 Anglesey, I am engaged in a review of the current records. I would be particularly pleased to receive any old Anglesey records which have been lying neglected in field notebooks, or which derive from specimens in collections. Anyone contributing such records or with information which leads me to useful historic material, will be welcome to receive a free copy of the North West Wales Lepidoptera Report. Volumes so far produced are 2001, 2002, and 2003, with 2004 in preparation (early 2006).

Julian Thompson and Andrew Graham would be similarly pleased to receive any records relating to Caernarvonshire and Merionethshire, respectively. In tandem with the Lepidoptera Report, Andrew Graham has for some years been constructing an impressive database program for the area covered by VCs 48, 49 and 52. The database draws on the currently available records (at present comprehensive for VC48 - partial for VCs 49 and 52) and gives flight histograms, dot distribution maps, and much other useful information for more than 1 300 taxa. Copies of the database CD are available from Andrew Graham; SAE appreciated.

John Harold, Hen Ardd, Carreg y Gath, Rhiwlas, Bangor, Gwynedd, LL57 4HD

THE MAITLAND EMMET BENHS RESEARCH FUND

In 2001 the family of the late Lt. Col. Maitland Emmet, a distinguished amateur microlepidopterist, made a generous donation to the Society’s Research Fund in his memory As a result the Society has renamed its Research Fund the Maitland Emmet BENHS Research Fund. The Society is very grateful to the Emmet family for their generosity.

The Society invites applications for grants, from the Maitland Emmet Research Fund, to be awarded in December 2006. Awards are open to both members and non-members of the BENHS and will be made to support research on non-marine arthropods, with reference to the British fauna, and with preference given to insects, arachnids, myriapods and isopods. Grants will be given for:

(a) the assistance of fieldwork on non-marine arthropods with relevance to their conservation,

(b) work leading to the production of identification guides and distribution lists, but not the cost of publishing such items.

Travel to examine museum collections and to consult taxonomic specialists would be included. The work and travel is not limited to the British Isles but must have a demonstrable relevance to the British arthropod fauna. Individual grants are unlikely to exceed £500.

Preference will be given to work with a clear final objective (e.g., leading to publication or the production of a habitat management plan). Work on leaf miners and gall forming insects should be submitted to the Society’s Professor Hering Memorial Research Fund.

Applicants should send seven copies, if possible, of their plan of work, the precise objectives, the amount for which an award is requested and a brief statement outlining their experience in this area of work, to Dr J. Muggleton, 17 Chantry Road, Wilton, Salisbury, Wiltshire SP2 0LT, as soon as possible and not later than 30 September 2006. Further information may be obtained from the same address (email: jmuggleton@aol.com).

THE PROFESSOR HERING MEMORIAL RESEARCH FUND

The British Entomological and Natural History Society announces that awards may be made from this Fund for the promotion of entomological research with particular emphasis on:

(a) leaf-miners

(b) Diptera, particularly Tephritidae and Agromyzidae

(c) Lepidoptera, particularly Microlepidoptera

(d) general entomology

in the above order of preference having regard to the suitability of applicants and the plan of work proposed.

Awards may be made to assist travelling and other expenses necessary for fieldwork, for the study of collections, for attendance at conferences, or, exceptionally, for the costs of publication of finished work. In total they are unlikely to exceed £1000 in the year 2006.

Applicants should send seven copies, if possible, of a statement of their qualifications, of their plan of work, and of the precise objectives and amount for which an award is sought, to Dr J. Muggleton, 17 Chantry Road, Wilton, Salisbury, Wiltshire SP2 0LT as soon as possible and not later than 30 September 2006.

Applications are also invited from persons wishing to borrow the Wild M3 Stereomicroscope and fibre optics illuminator bequeathed to the Fund by the late Edward Pelham-Clinton, 10th Duke of Newcastle. Loan of this equipment will be made for a period of up to six months in the first instance.

SMITHSONIAN INSTITUTION LIBRARIES

3 9088 90052 7300

BRITISH JOURNAL OF ENTOMOLOGY AND NATURAL HISTORY VOLUME 19, PART 1, APRIL 2006

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41

37

38

40

ARTICLES

A sawfly, Pristophora leucopus (Hellen), (Hymenoptera: Tenthredinidae) new to Britain.

K. J. Grearson

The modern bee and wasp assemblages (Hymenoptera: Aculeata) of Warwickshire’s calcareous quarries and spoilheaps, and the conservation issues facing them.

S. J. Falk

New county records of Heteroptera (Hemiptera) from Gloucestershire.

K. N. A. Alexander

On the identity of Pseudexechia parallela (Edwards, 1925) (Diptera: Mycetophilidae) and description of a new related species from Great Britain.

J. Klerandsen & P. J. Chandler

SHORT COMMUNICATIONS

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), the multi-coloured Asian ladybird, an etymological note.

J. Muggleton

Schreckensteinia festaliella (Hiibner) (Lepidoptera: Schreckensteiniidae) on an unusual food- plant in the Outer Hebrides.

K. P. Bland

Mortality of Orthoptera caused by mechanised mowing of grassland.

T. Gardiner

Exochomus quadripustulatus (L.) (Coleoptera: Coccinellidae) as a host of Dinocampus coccinellae (Schrank) (Hymenoptera: Braconidae).

P. R. M ABBOTT

PROCEEDINGS & TRANSACTIONS/SOCIETY NEWS

50 Results of the 2005 BENHS members’ questionnaire M. G. Telfer

REVIEW

6 The Empidoidea (Diptera) of Fennoscandia and Denmark. IV Genus Hilara. Fauna

Entomologica Scandinavica 40 by M. Chvala. A. Plant

ANNOUNCEMENTS

49 Status of The Moths and Butterflies of Great Britain and Ireland

64 Lepidoptera recording in northwest Wales (Merionethshire, Caernarvonshire and Anglesey).

J. Harold

CORRECTION

6 Re: Barrington, R.D.G. & White, M.C.

Editor

June 2006

ISSN 0952-7583

Yol. 19, Part 2

i

BRITISH JOURNAL OF

ENTOMOLOGY

AND NATURAL HISTORY

BRITISH JOURNAL OF ENTOMOLOGY AND NATURAL HISTORY Published by the British Entomological and Natural History Society and incorporating its Proceedings and Transactions

Editor: J. S. Badmin, Coppice Place, Perry Wood, Selling, nr Faversham, Kent ME13 9RB (Tel: 01227 752291) email: jbadmin@btinternet.com

Associate Editor: M. Wilson, Ph.D., F.R.E.S., F.L.S. Department of Biodiversity & Systematic Biology, National Museums & Galleries of Wales, Cardiff CF10 3NP. (Tel: 02920 573263) email: Mike.Wilson@nmgw.ac.uk

Editorial Committee:

D. J. L. Agassiz, M.A., Ph.D., F.R.E.S. R. D. G. Barrington, B.Sc.

P. J. Chandler, B.Sc., F.R.E.S.

B. Goater, B.Sc., M. I. Biol.

A. J. Halstead, M.Sc., F.R.E.S.

R. D. Hawkins, M.A.

P. J. Hodge

T. G. Howarth, B.E.M., F.R.E.S. I. F. G. McLean, Ph.D., F.R.E.S M. J. Simmons, M.Sc.

P. A. Sokoloff, M.Sc., C.Biol.,

M. I. Biol., F.R.E.S.

R. W. J. Uffen, M.Sc., F.R.E.S. B. K. West, B.Ed.

British Journal of Entomology and Natural History is published by the British Entomological and Natural History Society, Dinton Pastures Country Park, Davis Street, Hurst, Reading, Berkshire RG10 OTH, UK. Tel: 01189-321402. The Journal is distributed free to BENHS members.

© 2006 British Entomological and Natural History Society.

Typeset by Dobbie Typesetting Limited, Tavistock, Devon.

Printed in England by Henry Ling Ltd, Dorchester, Dorset.

BRITISH ENTOMOLOGICAL AND NATURAL HISTORY SOCIETY Registered charity number: 213149

Meetings of the Society are held regularly in London, at the rooms of the Royal Entomological Society, 41 Queen’s Gate, London SW7 and the well-known ANNUAL EXHIBITION is planned for Saturday 11 November 2006 at Imperial College, London SW7. Frequent Field Meetings are held at weekends in the summer. Visitors are welcome at all meetings. The current Programme Card can be obtained on application to the Secretary, J. Muggleton, at the address given below.

The Society maintains a library and invertebrate collections at its headquarters in Dinton Pastures, which are open to members on various advertised days each month, telephone 01189-321402 for the latest meeting news. The Society’s web site is: http://www.BENHS.org.uk

Applications for membership to the Membership Secretary: D. Young, 22 Wordsworth Close, Saxmundham, Suffolk IP 17 1WF. Tel: 01728 603568.

Subscriptions and changes of address to the Assistant Treasurer: R. D. Hawkins, 30D Meadowcroft Close, Horley, Surrey RH6 9EL.

Non-arrival of the Journal, faulty copies or other problems arising from distribution of the Journal or notices to the Distribution Secretary: D. Young, 22 Wordsworth Close, Saxmundham, Suffolk IP 17 1WF. Tel: 01728 603568.

Orders for books and back numbers of the Journal and Proceedings to the Sales Secretary: G.

Boyd, 91 Fullingdale Road, Northampton NN3 2PZ. Tel: 01604 410056.

General Enquiries to the Secretary: J. Muggleton, 17 Chantry Road, Wilton, Salisbury, Wiltshire SP2 0LT. Tel: 01722 741487. email: jmuggleton@aol.com

Society Website: www.benhs.org.uk for recent information on the Society’s meetings programme and general society details.

Cover photograph: Hyphoraia testudinaria (Geoff.) (Arctiidae), at mv light, Chichester, West Sussex, 30.V.2005, collected by Sarah Patton. Wingspan 50 mm. Photo: R. A. Jones.

NOTE: The Editor invites submission of photographs for black and white reproduction on the front covers of the journal. The subject matter is open, with an emphasis on aesthetic value rather than scientific novelty. Submissions can be in the form of colour or black and white prints or colour transparencies.

BR. J. ENT. NAT. HIST., 19: 2006

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THE ACULEATE HYMENOPTERA OF SHOTOVER HILL, OXFORDSHIRE

Ivan R. Wright1 and Steve J. Gregory2

1 Shot over Wildlife, 15, Blenheim Way, Horspath, Oxford, 0X33 1SB 2Northmoor Trust: Ecology Department, Little Wittenham, nr Abingdon, Oxon, 0X14 4RA

Abstract

Five years of new survey work on Shotover Hill (2000-2004) for aculeate Hymenoptera are presented and compared with newly collated historic records. This study has recorded 1 84 species, which when combined with the results of surveys in the 1980s, places the total of recently recorded aculeate species at 209. By comparison, in the years before 1939 an estimated total of 203 species had been accumulated. Sixty-two of the species that were recorded before 1939 have not been observed in recent years, yet the recent work (1980s-2004) has added 68 species (54 by this study). The nesting requirements of these ‘formerly recorded’ and ‘recently added’ species differ, and are shown to be consistent with the land use changes since the 1930s. Site quality scoring indices suggest that when compared with other UK sites, Shotover Hill remains an important site for wasps and bees. The all-time total of aculeate Hymenoptera for Shotover Hill is calculated to be 271 species.

Introduction

Like the many small areas of sandy soil in Oxfordshire, Shotover Hill developed as heath through being suitable only for rough grazing and pasture. In the 20th century, Shotover was spared the agricultural ‘improvement’ of many local heaths through its proximity to Oxford, gaining a measure of protection as an area of public enjoyment and academic study.

Much of the current interest in aculeate Hymenoptera on Shotover Hill stems from work in the early years of the 20th century by staff of Oxford University Museum of Natural History. By 1939, 141 species had been recorded for Shotover (Salzman, 1939), rising to 185 species by 1987 (Steel, 1984; M. E. Archer, pers. comm.). Subsequent work on the source material by the authors has shown that when common and widespread species are included from Salzman (1939), the overall total of Aculeata for Shotover Hill by the 1980s (before the addition of records presented in this study) is estimated to be 217 species.

A site with > 150 species of aculeate can be considered ‘Class V in Britain (Steele, 1984), especially for locations that are some distance from East Anglia and the south coast of Britain (M. E. Archer, pers. comm.). Clearly, Shotover Hill would rank as an important site on the basis of accumulated species, however 99 of these species were not re-recorded by the subsequent work of the 1980s (Steel, 1984; M. E. Archer pers. comm.). Since the 1930s, scrub encroachment on Shotover has considerably reduced the extent of open grassland, heath and bare soil, and without further survey work, an historic total of 185 species may not be a true reflection of the current status of the Aculeata on Shotover Hill.

Site Description

Shotover Hill (summit: 171m amsl at SP564063) is 3 km east of Oxford, and is a small sandstone plateau, 100m above the Thames Clay Vales and within the Midvale

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Heath and

acid grassland

Whitchurch Sand Portland Limestone Glacial 'Head'

Iv inline ridge Sand Kimmeridge Clay Ampthill Clay Open water

Aculeate

sampling point

Summit: 171m,

SP564063

Field boundary

SSSI boundary

Figure 1. A plan of Shotover Hill showing the principal aculeate sampling points, geology and areas of heath and acid grassland.

Ridge Natural Area (English Nature, 1997). Below the Cretaceous capping of Whitchurch Sand (Fig. 1) are strata of Portland Limestone and Kimmeridge Clay (Jurassic) which, at Shotover, includes associated beds of sand and coarse stone with a range of different textures. In places, the lower slopes are covered with shallow deposits of glacial clay or ‘head’. The land use is mixed and comprises heathland scrub, secondary woodland of various ages, pasture, large gardens and a little arable agriculture. The area represented by surveying in this study is about 250 hectares.

Much of the south west side of Shotover Hill is taken up with a public access Country Park, most of which is designated an SSSI (Fig. 1) for its flora and invertebrate fauna. Over recent years, the heathland on Shotover has been enlarged and although the area of Calluna heather is small (about 1.5 hectares), it is a relatively large expanse for the County of Oxfordshire.

On the basis of archive photographs spanning the 20th century (including many aerial photographs) and anecdotal evidence from local residents, it has been possible to interpret the changing habitats over this period. Until the middle years of the 20th century, Shotover was predominantly rough grazing and low-grade pasture with some areas of crop cultivation and isolated woodland. Then, between c. 1940 and c.1960, the area was considerably altered through various significant influences: the

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reduction of pasture-based farming, reduced rabbit grazing following the onset of myxomatosis, and a period of substantial disturbance by public and military vehicles.

Therefore, until about 1950 extensive areas of short vegetation and bare soil would have been present, and would have been of considerable benefit to ground nesting invertebrates. Since 1950, scrub vegetation has grown up and matured in places, creating extensive pioneer woodland with only isolated areas of mown or rabbit- grazed grassland.

Assessment of Historic Records

Appendix 12 of Steel (1984) provides a list of aculeate Hymenoptera recorded on Shotover Hill to that date: that is, Salzman (1939) plus the later work of O’Toole. However in returning to the original data, possible errors in the extraction of data from Salzman (1939) were highlighted, and this accounts for small differences between the species given here and those in Steel (1984). For this study, six species are discounted from Steel’s list: Ancistrocerus antilope (Panzer), Passaloecus gracilis (Curtis), Andrena bucephala (Stephens), A. nigroaenea (Kirby), Bombus ruder atus (F.) and B. rupestris (F.), and 1 1 species are added: Arachnospila anceps (Wesmael), Ancistrocerus oviventris (Wesmael), A. trifasciatus (Muller), Crabro peltarius (Schreber), Mimesa equestris (F.), Passaloecus monilicornis Dahlbom, Andrena pilipes (F.), A. trimmerana (Kirby), Anthidium manicatum (L.), Nomada flavoguttata (Kirby) and Bombus sylvestris (Lepeletier). This adjustment increases Steel’s accumulated total for that time (1984) from 174 to 179 species.

Incorporating the above adjustment, Salzman ( 1939) lists 141 species of aculeate as being specifically associated with ‘Shotover’, and a further 62 species are listed as widespread or ‘common round Oxford’. Many of these common species would have been present on Shotover Hill, and probably recorded there, but were not specifically listed as such. The number of species actually recorded on Shotover in the early 20th century must, therefore, be between the minimum of 141 and a maximum of 203 (Table 1), and introduces some uncertainty into the interpretation of any subsequent changes in the aculeate fauna.

Table 1. Summary of previous aculeate Hymenoptera recording on Shotover Hill.

Historic records	Number of species based on ‘1939 minimum’	Species ‘common round Oxford’ (1930s)	Number of species based on 4939 maximum’
Species recorded before 1939	141	+ 62	203
(Salzman, 1939) of which Species remaining unrecorded by the 1980s	99	+ 32	131
Species refound in the 1980s by O’Toole and Archer	42	+ 30	72
Species added in the 1980s by O’Toole and Archer	44		14
Total species recorded by the 1980s	185	+ 32	217

‘1939 minimum’ includes species specifically recorded for ‘Shotover’ in Salzman (1939)

‘1939 maximum’ includes all species listed as either for ‘Shotover’ or ‘common round Oxford’.

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® Specifically recorded Shotover records Recorded between

for Shotover Hill + 'common' species 1985 and 2004 before 1939 before 1939 (209 species)

(141 species) (203 species)

Figure 2. Status profiles of aculeate Hymenoptera records showing the effect of including implicit ‘common’ species from earlier records. National statuses after Falk (1991) and Ball (1997).

In the early 1980s, Mr Chris O’Toole (Department of Entomology, Oxford University) made two visits to survey Aculeata on Shotover Hill, and Dr Michael Archer (BWARS) made survey visits in 1985 and 1987. These visits recorded a total of 86 species, of which 42 had been previously noted for Shotover Hill by Salzman (1939), and 44 were formally new records for Shotover (Table 1). However, when considering those species that Salzman notes as ‘common round Oxford’, these statistics are altered to 72 species re-recorded and 14 added. Consequently, the all- time total of Aculeata on Shotover Hill by 1987 was formally raised to 185, or when ‘common’ species are added, to an estimated total of 217 (Table 1).

Table 1 shows that the exclusion of common aculeate species would greatly affect a comparison between current and previous work. Therefore it is necessary to investigate some feature of the recorded species to justify either exclusion of the implicit ‘common’ species (i.e. ‘1939 minimum’ in Table 1) or their inclusion (i.e. ‘1939 maximum’). It is proposed here that the national statuses for Aculeata (Falk, 1991; Ball, 1997) would be a suitable indicator, even though the conservation status of some species will have changed since the 1930s.

Figure 2 shows that there is a very marked similarity between the status profiles of early and recent work, but only when the ‘common’ species are included. Although some of the ‘common’ species may not have been recorded on Shotover in the 1930s, the inclusion of all ‘common’ species from Salzman (1939) would clearly provide the most relevant comparisons with recent work. It is unlikely that changes in status over time would greatly affect this conclusion, and all ‘common’ species (1939 maximum) are assumed to have been on Shotover in the 1930s for the purposes of analysis herein.

Survey Methodology (2000-2004)

Specimens were collected over the five years from 2000 to 2004, and sampled by water trap. Malaise trap and by hand net. Pitfall traps yielded most of the ants and a few other Hymenoptera.

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Most of the trapping activity was concentrated at the points shown in Fig. 1. Although the distribution of sampling is reasonably representative of the area, it was not possible to give the same amount of attention to all locations, and not all locations were sampled using the same methods. Some of the sampling was within the public park where passive traps cannot be left unattended for more than a day or two without risk of disturbance. However, some of the areas of heavy public usage coincide with the heathland habitat where many of the scarcer Aculeata would be expected. In these areas there is, therefore, a bias towards netting by hand. In light of these limitations, no attempt has been made to analyse the species profiles from the areas of different soil or habitat within the site.

Results

In this study 184 species of aculeate Hymenoptera were recorded during the period 2000-2004 (Appendix 1), of which 54 are new records for Shotover Hill. Table 2 gives a summary of these species by family/sub-family (BWARS, 2005), and Table 3 shows a summary of their national statuses (Falk, 1991; Ball, 1997), together with more recent ratings derived from national records for the period 1970 to 2004 (Archer, 2004a, 2004b, 2005; BWARS, 2003). The status of five species were 'unknown’ to Ball (1997) but have been classified since by Archer (2004a & b) as either ‘universal’ or ‘widespread’. For completeness these five species have been given retrospective equivalent statuses, however, this does not affect the analysis that follows.

When the records from this study are supplemented by the work of C. O’Toole and M. E. Archer in the 1980s, a further 25 species can be included in the analysis as representative of recent recording (Appendix 2). From the early 1980s until 2004, 209 species have been recorded, of which 1 16 are formally new records for Shotover Hill. However, 48 of these 116 species are amongst those that Salzman (1939) considered ‘common round Oxford’, leaving 68 species as entirely new records for Shotover.

Table 2. Tabulation by family/subfamily of species recorded in this study (2000-2004).

	Family	No. of species
Ants	Formicidae	8
Wasps	Chrysididae	6
	Tiphiidae	2
	Mutillidae	1
	Pompilidae	13
	Eumeninae	4
	Vespinae	4
	Crabonidae	47
Bees	Colletinae	8
	Andreninae	24
	Halictinae	28
	Megachilinae	11
	Anthophorinae	17
	Apinae	11
All species	Total	184

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Table 3. Tabulation by national status of species recorded in this study (2000-2004).

Status Falk (1991) Ball (1997)	No.	Status Archer (2004a&b, 2005) BWARS (2003)	No.
Common	89	Universal	92
Local	65	Widespread	57
Nationally Scarce b	14	Restricted	11
Nationally Scarce a	13	Scarce	17
Red Data Book 3	2	Rare	6
Red Data Book 2 Red Data Book 1	1 0	Very Rare	1
Totals	184		184

Table 4. Quality scoring indices for the recent surveys (1980s-2004) and pre-1939 on Shotover Hill, together with quality indices for other UK sites of comparative area.
	Area	No. of	Quality Score	Aerial Nester Frequency	Parasitic Load
Site	(ha)	Species	Wasps	Bees	Wasps	Bees
Chafford Hundred, Essex1	137	218	5.96	20.3	22.3	19.5	26.4
Ambersham Common, West Sussex2	212	219	4.53	38.5	23.3	16.1	22.3
Iping Common, West Sussex2	172	219	4.51	42.4	21.0	12.4	23.5
Sherwood Forest, Nottingham- shire3	390	100	2.96	47.6	19.4	17.6	25.0
Shotover 1980s-20044	250	209	2.89	51.3	27.4	23.1	32.3
Shotover pre-19394	-250	203	-	44.6	23.4	29.6	29.6
Crow Wood, Yorkshire3	152	105	2.53	20.4	9.4	16.9	28.9
Blaxton Common, Yorkshire3	150	109	1.85	43.1	13.9	15.0	26.5
Risby Warren, Lincolnshire3	170	63	1.81	12.5	8.3	17.2	29.4
Skipworth Common, Yorkshire3	312	69	1.57	42.4	30.0	13.2	35.5

‘O’Toole (1998)

2Archer & Edwards (2002) 3 Archer & Burn (1995) 4This study

Of the species listed for the early 20th century by Salzman (1939) 62 remain unrecorded by the recent surveys (Appendix 3). Including all surveys, the total number of species so far recorded on Shotover Hill will be close to the estimated maximum total of 271: about half of the national fauna of Aculeata.

Quality scoring indices

Following work by Archer & Burn (1995) to derive indices that describe the aculeate fauna of a site, quality scores have been calculated for Shotover Hill in a similar manner. These are the Quality Score, Aerial Nester Frequency and Parasitic Load, and permit objective comparisons with other UK sites for which similar work

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has been presented. The Quality Score used here is the total score after assigning each recorded species with a value relating to its national status (but not its abundance): e.g., common = 1, very rare = 32, (Archer & Burn, 1995), divided by the total number of species in the survey. Thus the calculated index includes a measure of the proportion of scarcer species at each site. For the most recent survey work on Shotover Hill (2000-2004) the Quality Score is 2.72, which increases to 2.89 when combined with the work in the 1980s by O’Toole and Archer.

Aerial Nester Frequency expresses the number of Aculeata that nest above ground as a proportion of the total soil and aerial nest builders, and Parasitic Load expresses the number of parasitoids and cleptoparastites as a proportion of the total host and parasite species. These indices are calculated for bees and wasps separately. Table 4 shows published indices for eight UK sites that are notable for their aculeate Hymenoptera (Archer & Burn, 1995; O’Toole, 1998; Archer & Edwards, 2002). As Quality Scores are partly influenced by the area of a site (Archer & Burn, 1995), only those sites in the range 100-400 ha were chosen, being within 150 ha of the area surveyed in this study. Table 4 lists the chosen sites in order of Quality Score, which places Shotover Hill centrally among the other notable sites.

It would not be relevant to calculate the site Quality Score for the historic data from Shotover Hill, as the statuses would not be accurate for the early 20th century, and comparison could be misleading. However, Aerial Nester Frequency and Parasitic Load can be legitimately derived for both historic and recent data, and these are shown in Table 4.

Discussion

Rare and notable species

Considering the influences of changing habitat, and possibly climate, it is to be expected that the status of mobile species such as Hymenoptera will change over time, and Table 3 should be viewed with this in mind. Of the three 'Red Data Book’ (RDB) species, the cleptoparasitic bee Sphecodes niger (von Hagens) is now listed as ‘very rare’ in Britain (Archer, 2005). However, the ‘Bee Wolf wasp, Philanthus triangulum (F.) (RDB2 but provisionally RDB4) and cleptoparasitic bee Nomada lathburiana (Kirby) (RDB3) have become much less scarce and are now considered widespread (Archer 2004a & b). Table 3 shows that 30 (16%) of the species recorded by this study are either ‘Nationally Scarce’ or ‘RDB’, but this is not an especially high proportion compared with other published studies (e.g. O’Toole, 1998; Archer & Edwards, 2002). However, it should be noted that it is easier to observe a species becoming more common by recording its presence, than it is to suspect that a species is becoming scarcer by eventually noting its absence.

Several are new records for this far north in the region: for example the solitary wasp Astata boops (Schrank) and its associated cleptoparasite Hedychridium roseum (Rossius). Were it not for the paucity of recent recording of Aculeata in the region, these new records could be indicators of northward migration.

Comparison with historic records

Reworking past records from Shotover Hill has shown that in the early 20th century the sandy heath and pasture of the hill supported a remarkable diversity of aculeate Hymenoptera. Notwithstanding the uncertainty of including implicit common species, a total of about 200 species would rank as a very good habitat by current standards. In comparison, this and recent studies (1980s-2004) have

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Table 5. National status and nesting requirements of ‘Formerly recorded’, ‘Recently added’ and ‘Continuously present’ species on Shotover Hill.

	‘Formerly recorded’ Species not recorded since 1939. %	‘Recently added’ Species newly added 1980s-2004. %	‘Continuously present’ Species recorded in both early and recent surveys. %
Common	19	36	53
Local	40	41	33
Nationally Scarce b	18	6	10
Nationally Scarce a	8	13	4
RDB	15	4	0
	No. (%)	No. (%)	No. (%)
Soil nesters	30 (48)	26 (38)	65 (46)
Aerial nesters	10 (16)	23 (34)	37 (26)
Total species	62	68	141

recorded 209 species, which includes 68 that were either not present or not recorded on Shotover in the 1930s. Clearly some of these species went unrecorded, such as the common ant Myrmica ruginodis Nylander, but 68 species seems to be too many to have gone unnoticed by the experienced collectors of Oxford University in those early years. Considering the 62 species that have not been observed since 1939, the habitat requirements of these ‘formerly recorded’ and ‘recently added’ groups of aculeate species warrants investigation.

Table 5 shows the distribution of national statuses within the ‘formerly recorded’, ‘recently added’ and ‘continuously present’ groups of species. Although some species may have been present but not recorded in the various surveys, and current statuses may not be representative of the 1930s, the large number of species involved would suggest that most of the differences are due either to habitat change, or species becoming generally more scarce or common in the region.

As might be expected, there is a high proportion of the more ubiquitous common species (53%) recorded continuously through both surveys. Of the scarce and possibly more habitat specific species, just 14% (all Nationally Scarce and no RDBs) have survived on Shotover from the early 20th century to the present day (Table 5). In contrast to this, 41% and 23% of ‘formerly recorded’ and ‘recently added’ species respectively are Nationally Scarce or Red Data Book. (Separate comparison of the ‘formerly recorded’ and ‘recently added’ groups also mitigates the earlier decision to include the implicit ‘common’ species.)

Of particular interest is the proportion of ‘formerly recorded’ and ‘recently added’ species that build their nests specifically either in the soil or in aerial cavities (e.g., hollow plants stems and holes in dead wood). In Table 5 it can be seen that there are more soil nesters in the ‘formerly recorded’ group of species (30) than in the ‘recently added’ group (26), yet many more aerial nesters have been ‘recently added’ (23) than are in the ‘formerly recorded’ group (10). The change over time from soil to aerial nesting is statistically significant at the 5% level (d.f. = 1, %2 = 4.54, p<0.05), and is entirely consistent with the habitat changes over the period. In the early 20th century, Shotover had much more pasture and patches of sunlit soil, especially on

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the warmer south side of the hill. In contrast to this, much of the obsolete pasture and rabbit-grazed grassland has now succeeded to woody scrub, and bare soil is reduced to paths, areas of conservation effort and public disturbance. When wasps and bees are considered separately, Table 4 shows a very similar increase (~ 16%) in Aerial Nester Frequency for both groups since the 1930s.

Comparing the pre-1939 and post 1980s surveys, Parasitic Load for bees rose a little over the interceding years from 29.6 to 32.3 ( + 9%), and for wasps fell from 29.6 to 23.1 ( — 22%). However, these changes are not statistically significant. When compared with the Parasitic Load values given for the other eight UK sites (Table 4), Shotover appears to have a higher proportion of parasitic species than most of the other listed sites.

Conclusion

The 184 species recorded by this study show that, on this basis alone, Shotover Hill retains a good diversity of aculeate Hymenoptera. An area of 250 hectares in the south of England, which includes a range of heath and scrub woodland habitats, would be expected to yield a reasonable diversity of Aculeata. However, Shotover Hill is some distance from the particularly diverse southern coastal sites and is therefore notable in supporting such a diversity of Aculeata in the south midlands of Britain. Furthermore, several of the species recorded in this study are close to the north-western limit of their UK and European ranges.

The recording of aculeate Hymenoptera on Shotover Hill in this study, and over the past 100 years, has shown that the overall species diversity has remained fairly constant at just over 200 species, even though the land use and vegetation have changed. However, as pasture and short heathy scrub have been replaced, in part, by successional woodland, it appears that the profile of aculeate species has changed in a manner that this succession would suggest. When comparing those species that have not been recorded since 1939, for whatever reason, with those that have been recorded more recently, there has been a small but consistent shift from soil to aerial nesting in both wasps and bees. Measures could be taken to increase the range of bare soil habitats, and thereby avoid a possible trend towards fewer soil nesting species. Nevertheless, this study of aculeate Hymenoptera has shown that Shotover Hill remains an important habitat for this specialised group of invertebrates.

Acknowledgements

The authors would like to express their thanks to the following for permission of access and recording for the purposes of survey work: Lt. Col Sir John Miller of Shotover House, Oxford City Council, Mrs D. Stogdale of Monk’s Farm, Shotover and Mr B. Walker of Manor Farm, Horspath. We also express our thanks for the assistance of the Oxfordshire Biological Records Centre (now the Thames Valley Environmental Records Centre), the Northmoor Trust Ecology Department (Little Wittenham, Oxfordshire) and the Department of Entomology at the Oxford University Museum of Natural History. Also thanks to Mr C. O’Toole and Dr M. E. Archer for additional information and unpublished data.

References

Archer, M. E. 2004a. Archer’s status values for the solitary wasps. BWARS Newsletter Spring , 2004: 22-24.

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Archer, M. E. 2004b. Archer’s status values for the solitary bees. BWARS Newsletter Autumn , 2004: 31-33.

Archer, M. E. 2005. Archer’s status values for the solitary wasps and bee species. BWARS Newsletter Spring , 2005: 17-19.

Archer, M. E. & Burn, J. T. 1995. The aculeate wasps and bees of Crow Wood, Finningley in Watsonian Yorkshire, with the introduction of a new national quality scoring system. British Journal of Entomology and Natural History 8: 49-59.

Archer, M. E. & Edwards, M. 2002. The aculeate Hymenoptera of Ambersham and Iping (with Stedham) Commons in West Sussex, including statistical procedures for estimating species richness. British Journal of Entomology and Natural History 18: 91-103.

Ball, S. G. 1997. RECORDER 3.3: An environmental recording package for local recording centres. JNCC, Peterborough, UK.

BWARS 2003. Bees, Wasps and Ants Recording Scheme Web Site, http://www.bwars.com. Aculeate Species Listing file, \ASL_03-04-03_tst.txt’

BWARS 2005. Check list of British Hymenoptera Aculeata. In Bees, Wasps and Ants Recording Scheme Members’ Handbook. CEH, Monk’s Wood, UK.

English Nature 1997. Midvale Ridge Natural Area. English Nature, Newbury.

Falk, S. J. 1991. A review of the scarce and threatened bees, wasps and ants of Great Britain. Research and Survey in Nature Conservation. No. 35. Nature Conservancy Council, Peterborough.

O’Toole, C. 1998. The Chafford Hundred experience: surveying aculeate Hymenoptera in a large, post-industrial building development. In: Jones, R. & Munn, P. (Eds.) Habitat management for wild bees and wasps. International Bee Research Association, Cardiff.

Salzman, L. P. (Ed.) 1939. Victoria County History of Oxfordshire. University of London Institute of Historical Research. Vol. 1. Oxford University Press.

Steel, D. 1984. Shotover: The Natural History of a Royal Forest. Pisces Publications, Oxford.

Appendix 1

Aculeate Hymenoptera (184 species) recorded on Shotover Hill, 2000-2004. National statuses (BWARS, 2003; Archer, 2004a, 2004b & 2005) as Universal (U), Widespread (W), Restricted (RE), Scarce (S), Rare (R), Very rare (VR) and (new) indicates a new record for Shotover Hill by this study.

Chrysididae: Hedychridium roseum (Rossius) (S, new), H. ardens (Latreille in Coquebert) (U), Trichrysis cyanea (L.) (W), Chrysis angustula Schenck (W, new), C. ignita (L.) (U), C. impressa Schenck (U, new).

Tiphiidae: Tiphia femorata F. (S, new), T. minuta Vander Linden (W).

Mutillidae: Myrmosa atra Panzer (W).

Formicidae: Myrmica lobicornis Nylander (U, new), M. rubra L. (U), M. ruginodis Nylander (U, new), M. scabrinodis Nylander (U, new), Formica fusca L. (U), Lasius flavus (F.) (U), L. brunneus (Latreille) (RE, new), L. niger (L.) (U).

Pompilidae: Priocnemis perturbator (Harris) (U), P. coriacea Dahlbom (R), P. parvula Dahlbom (U), P. exaltata (F.) (U, new), P. fennica Haupt (W, new), P. hyalinata (F.) (S, new), P. gracilis Haupt (S), P. schioedtei Haupt (U), Dipogon subintermedius (Magretti) (U, new), Caliadurgus fasciatellus (Spinola) (S, new), Arachnospila anceps (Wesmael) (U), Evagetes crassicornis (Shuckard) (U), Anoplius nigerrimus (Scopoli) (U, new).

Vespidae: Gymnomerus laevipes (Shuckard) (S), Symmorphus bifasciatus (L.) (U), Ancistrocerus gazella (Panzer) (W, new), A. trifasciatus (Muller) (U). Dolichovespula sylveslris (Scopoli) (U), D. media (Retzius) (W, new), Vespula vulgaris (L.) (U), V. germanica (F.) (U).

Crabronidae: Astata boops (Schrank) (RE, new), Tachysphex pompiliformis (Panzer) (U), Trypoxylon attenuatum Smith, F. (U), T. clavicerum Lepeletier &

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Serville (W, new), T. medium de Beaumont (U, new), Crabro peltarius (Schreber) (U), Crossocerus ovalis Lepetelier & Brulle (U, new), C. pusillus Lepetelier & Brulle (U), C. cetratus (Shuckard) (W, new), C. megacephalus (Rossius) (U), C. podagricus (Vander Linden) (U), C. quadrimaculatus (F.) (W), C. dimidiatus (F.) (U), Ectemnius cavifrons (Thomson) (U), E. continuus (F.) (U), E. lituratus (Panzer) (RE, new), E. cephalotes (Olivier) (W, new), Rhopalum clavipes (L.) (U), Lindenius panzeri (Vander Linden) (RE, new), L. albilabris (F.) (U), Entomognathus brevis (Vander Linden) (W), Oxybelus uniglumis (L.) (U), Mimesa equestris (F.) (U), Mimumesa dahlbomi (Wesmael) (W, new), Psenulus pallipes (Panzer) (W), P. concolor (Dahlbom) (W, new), P. schencki (Tournier) (R, new), Pemphredon lugubris (F.) (U), P. inornata Say (U), P. lethifera (Shuckard) (U), P. morio Vander Linden (S, new), Passaloecus corniger Shuckard (W), P. insignis (Vander Linden) (W), P. gracilis (Curtis) (W, new), P. singularis Dahlbom (W, new), Spilomena curruca (Dahlbom) (W, new), Mellinus arvensis (L.) (U), Didineis lunicornis (F.) (S, new), Nysson spinosus (Forster) (U), N. trimaculatus (Rossius) (W), N. dimidiatus Jurine (S), Gorytes quadrifasciatus (F.) (W), Harpactus tumidus (Panzer) (U), Argogorytes mystaceus (L.) (U), Philanthus triangulum (F.) (W, new), Cerceris rybyensis (L.) (RE), C. arenaria (L.) (W).

Colletinae: Colletes succinctus (L.) (U, new), C. similis Schenck (W, new), C. daviesanus Smith, F. (U, new), Hylaeus cornutus Curtis (S), H. annularis (Kirby) (RE, new), H. communis Nylander (W), H. hyalinatus Smith, F. (W), H. confusus Nylander (U, new).

Andreninae: Andrena haemorrhoa (F.) (U), A. flavipes Panzer (RE), A. nitida, (Muller) (W), A. nigroaenea (Kirby) (U), A. bicolor F. (U), A. scotica Perkins (U), A. fucata Smith, F. (U), A. helvola (L.) (W), A. fulva (Muller in Allioni) (U), A. clarkella (Kirby) (U), A. apicata Smith, F. (S), A. fuscipes (Kirby) (U, new), A. denticulata (Kirby) (U), A. barbilabris (Kirby) (U), A. humilis Imhoff (S), A. chrysosceles (Kirby) (W), A. labiata F. (S), A. semilaevis Perez (U), A. falsifica Perkins (R, new), A. minutula (Kirby) (U), A. subopaca Nylander (U), A. ovatula (Kirby) (W), A. wilkella (Kirby) (U, new), A. dorsata (Kirby) (W).

Halictinae: Halictus rubicundus (Christ) (U), H. tumulorum (L.) (U), Lasioglossum leucozonium (Schrank) (W), L. quadrinotatum (Kirby) (R), L. lativentre (Schenck) (W), L. calceatum (Scopoli) (U), L. albipes (F.) (U), L. malachurum (Kirby) (RE, new), L. pauxillum (Schenck) (RE, new), L. fulvicorne (Kirby) (W), L. villosulum (Kirby) (U), L. punctatissimum (Schenck) (W), L. minutissimum (Kirby) (W), L. parvulum (Schenck) (W), L. rufitarse (Zetterstedt) (W, new), L. smeathmanellum (Kirby) (U), L. morio (F.) (W), L. leucopus (Kirby) (U), Sphecodes gibbus (L.) (W), S. reticulatus Thomson (S, new), S. monilicornis (Kirby) (U), S. pellucidus Smith, F. (W), S.