Distribution, status and ecology of the Ant-lion Euroleon nostras in England 1997

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69 INVESTIGATIONS INTO THE DISTRIBUTION, STATUS AND ECOLOGY OF T H E ANT-LION EUROLEON NOSTRAS (GEOFFROY IN FOURCROY, 1785) (NEUROPTERA: M Y R M E L E O N T I D A E ) IN ENGLAND DÜRING 1997 COLIN W. PLANT The first confirmed British record of an ant-lion came on 5 September 1931 at Gorleston, when a Single male was taken " . . . clinging to a paling about thirty inches from the ground, and in an apparently torpid condition" by C. G. Doughty (Doughty, 1931; Killington, 1932). The insect was identified as Myrmeleon formicarius by Claude Morley who, in an editorial footnote to Doughty (op. cit.) stated that the area had been practically unworked for over a hundred years and did not rule out the possibility of the species being native. Recent investigations by Howard Mendel at Ipswich Museum turned up Doughty's specimen in that institution (Morley Collection, number R. 1953-22) and this proved to be Euroleon nostras rather than Myrmeleon formicarius. It is hard to comprehend the original misidentification of a species with darkly spotted wings with one whose wings are hyaline; the names formicarum and formicarius do not appear in the synonymy of E. nostras listed in Aspöck et al„ (1980) though Mendel (1996), in considering this particular problem disagrees. The Suffolk find was of such interest that a lengthy report appeared in a local newspaper in which Morley described the possibility of the insect having been "merely blown across the intervening seventy miles of sea" from the Continent as " ... an extremely improbable contingency" (Morley, 1931). Killington (1932) considered that "While the species can not be placed on the British list on the strength of this capture [referring to the Suffolk specimen] it is quite possible it will yet be found to breed here and entomologists living in sandy districts should keep an eye open for pits made by the larvae". On 21st July 1994, Hilary and Geoff Welch found two adult Euroleon nostras on the road to the Minsmere RSPB reserve, adjacent to the toilet block at grid reference TM471671); the discovery was reported by Mendel (1996) who made the initial identification (subsequently confirmed by myself). One specimen was retained and the other, which interestingly had a "kinked abdomen" deformity, was released. The presence of this deformity is strongly suggestive of difficulties in eclosion which, in its turn, is indicative of a locally bred specimen. A third specimen was subsequently found on 31sl August 1994 by D. Fairhurst - squashed on the floor of the toilet block. The toilets are reputed to be cleaned daily and there is no reason to suppose that this is not so. Not unnaturally, news of this discovery spread rapidly through the entomological Community and it soon emerged, via Tony Irwin at the Castle Museum, Norwich that there was another record of this species - Mr T. R. Mitchell recorded a specimen of E. nostras that flew into a house in Corton, near Lowestoft (grid reference TM5496) on 29lh August 1988. He photographed the insect and released it and Dr Irwin subsequently made the identification from these pictures. This information has not been published. It is not at all clear why nobody encountered the distinctive larval pits for a

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further two years, though it may simply have been a lack of precise knowledge about where to look. However, between and 9th August 1996, Mike Edwards was fortunate enough to discover some whilst surveying insects at the Minsmere RSPB Reserve. Subsequent searches revealed numbers in excess of one thousand and on the second day no less than twelve newly emerged adults were observed drying their wings in the vicinity (Cottle et al., 1997). To complete the account, following the publication of the first proven breeding record for E. nostras (Cottle et al., 1996), Mr W. R. B. Hynd informed me that on 6 June 1992 he had encountered larval pits at a point on the coast between Southwold and Dunwich. After discussion with Mr Hynd, it was agreed that this locality must have been the woodland around Dunwich Friary at TM4770. Unfortunately there appears to be no information on the number of pits present at that time; thorough searching in 1997 did not reveal any at all. It was now clear that Euroleon nostras was a resident British insect in 1996. Further, it was fairly certain that a breeding population had been overlooked at Minsmere since at least 1994, possibly since 1931 and even, perhaps, since 1781. There was a clear need for further investigation and! consequently, the present survey was commenced. The project is subject tรถ joint funding by English Nature, the Royal Society for the Protection of Birds and by the author of this report. Active searching of potentially suitable areas for larval pits The eastern boundary of the principal search area is the mean high tide line on the east coast of England. At its southern end, the boundary passes along the mean high tide line of the River Deben and continues along this water course to the bridge in the village of Ufford (grid reference TM300519), whence it follows the minor road north to join the AI2 trank road at TM306545). From here it follows the centre of the A12 road through Saxmundham and Yoxford to Blyburgh. From Blyburgh, the boundary follows to A145 Road to Beccles then turns eastwards along the centre of the AI46 road towards Lowestoft turning northwards again before reaching this town to follow the centre of the AI 117 to Great Yarmouth, terminating in the town centre. Definition and assessment of breeding colonies The positions of accumulations of larval pits were marked on maps and the number of pits was recorded against date for each "colony". Because ant-lions frequently vacate pits and build new ones, it could not be automatically assumed that the presence of a pit was evidence of the presence of a larva. Two different methods were employed to assess the percentage of larval pits that were occupied: 1

The larva was extracted and examined. This was the favoured method since it also permitted an assessment of the developmental stage and allowed measurements to be taken; 2 The surface of the ground was smoothed over a known number of pits during the morning and revisited after three hours to see how many new pits had been excavated. Both methods yielded more or less identical results in terms of the percentage

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of pits occupied. A third method, dropping an ant into a pit to see if the larva was active, was not an efficient technique for two reasons: first, the shadow cast over the pit may have caused some larvae to retreat before the food was offered, and second, it proved extremely difficult to deliver a live and wriggling ant directly into the centre of a larval pit! The traditional method of extracting larvae from pits is to use a trowel; this proved almost totally unsuccessful in Suffolk, and the position of many pits on small ledges on the root plates of fallen trees made this impractical in any case. Rather more effective where the terrain permitted was the use of a large diameter metal kitchen sieve, scooped swiftly under the pit. The collected sand was then very carefully sieved and the larva, when present, was easily located amongst the residue of small stones and previous meals. Another method, favoured by the author, is to plunge a wide-mouthed glass specimen tube vertically downwards, very rapidly, over the very centre of the pit to a depth of at least two inches. The plug of sand in the tube is then gently emptied over the palm of the hand until the larva is found (see plate 6). Some success was also had by teasing the pit base with a twig so that the larva was stimulated into grasping it.

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Distribution of ant-lion colonies located Based on the information currently available from this survey, Euroleon nostras is, and possibly always has been, restricted in England to areas of sand-based geology in the Vice County of East Suffolk (the Suffolk Sandlings). The insect is apparently confined, in 1997, to 42 discrete aggregations of varying size, situated in 17 monads (1 Km x 1 Km squares) of the Ordnance Survey National Grid. The name, location and maximum recorded size in 1997 of each of the 42 colonies are summarised in Table 1. Colony Code A B(l)-B(4) C D(l) D(2) D(3) ECl) E(2) F(l) F(2) G(l) G(2) G(3) G(4) G(5) G(6) G(7) G(8) G(9) G(10) G(ll) G(12) G(13) G(14) G(15) G(16) H

Site name (OS map)

Grid reference

B laxhall Common J Corporate Belts 1 Hangman's New Wood Whin Hill 1 Whin Hill 2 Whin Hill 3 Sheepwash Lane 1 Sheepwash Lane 2 Scotshall Coverts 1 Scotshall Coverts 2 Westleton Walks 1 Westleton Walks 2 Westleton Walks 3 Westleton Walks 4 Westleton Walks 5 Westleton Walks 6 Westleton Walks 7 Westleton Walks 8 Westleton Walks 9 Westleton Walks 10 Westleton Walks 11 Westleton Walks 12 Westleton Walks 13 Westleton Walks 14 Westleton Walks 15 Westleton Walks 16 Minsmere toilet bank

TM384562 TM465611 TM453668 TM4667 TM4667 TM4667 TM4667 TM4667 TM4667 TM4667 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4668 TM4767

Table 1: Breeding colonies of Euroleon nostras in England

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Max. no. of larval pits 1997 13 450 clO 59 4 106 122 29 28 18 104 18 89 47 18 7 66 20 20 37 37 56 17 37 6 11 68


ANT-LION IN ENGLAND

I J K L M N P Q R S T U V W X Y Z

Minsmere Sand Martin site Dunwich Heath footpath Dunwich Heath Dunes Dunwich Heath North Dun wich Heath West (1) Dunwich Heath West (2) Dunwich Heath (other areas) Mill Road Triangie Westleton Pit 1 Westleton Pit 2 Dun wich Forest verge Toby's Walks Walberswick Common Minsmere Gravel Pit Mumbury Hills Sutton Hoo Dunwich Friary

TM4767

73

20 123

TM475679471680 TM478676 TM475688 TM469681 TM469863 TM4768

53 50 700 30 uncounted

TM4568 TM4468 TM4468 TM4571 TM4474 TM4875 TM449669 TM449677 TM289489 TM4770

12 112 56 1 c.250 47 c20 clOO zero* zero**

* The Sutton Hoo colony was reported by a member of the public but surveyors could not locate any pits during 1997 ** The Dunwich Friary colony held an unspecified number of individuals in June 1992 but in spite of searching no pits or larvae could be found in 1997. Locating adults Adults were looked for by direct searching, by beating or otherwise disturbing Vegetation and by examining spiders webs. Local lepidopterists were also alerted to the possibility of adults entering moth-traps. Like all true Neuroptera, ant-lions have a closed gut so that undigested food is not voided. This matenal is retained in the gut throughout pupation and is voided, on eclosion of the adult, as a solid pellet. This pellet has a distinctive and constant colour, shape and size which renders it an identifiable object. Sieving sand within colonies after all the larvae had pupated and all evidence of larval pits had been removed by environmental influences was very effective in revealing these faecal pellets to provide evidence that adults had emerged. Suffolk Euroleon nostras adults emerge around the end of July and through August, probably in tune with climatic conditions. Eggs are laid soon after emergence and the larvae hatch then feed rapidly in favourable conditions undergoing their first moult (to second instar) by September. They over-winter as second instar larvae in the Company of third instar larvae from the previous year. At the onset of warmer weather in the spring they commence feeding again and in June they moult for the second time to become third instar larvae. They then pass their second winter in this third and final instar, feeding again in the following spring forming a cocoon and entering the pre-pupal stage in

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late May. Adults emerge in July and August, probably over a protracted period.

Numbers of adults There were a minimum of 3071 larval pits at any one point in time in 1997 prior to the emergence of adults in that year. This figure is the sum of the maximum counts made at each of the 42 larval aggregations between April and July inclusive. Because ant-lions frequently vacate pits and build new ones, it could not be automatically assumed that the presence of a pit was evidence'of the presence of a larva. It was thus necessary to investigate the rate of occupancy amongst the observed pits; the methodology has been described above. It was discovered that 90% of larval pits were occupied at any one time. Cottle et al (1996) reported an estimated figure in excess of 3000 pits between 6 and 9 August 1996. Assuming a 90% pit occupancy, that represents 2700 larvae. This compares favourably with the observed figure of 2764 larvae in the period from April to July 1997, suggesting minimal winter-mortality.

Feeding behaviour Euroleon larvae repose horizontally to one side of the pit, level with its lowest point or just slightly above it, and attack prey from the side. They do not seem to rest vertically at the base of the pit with j a w s upwards as popularly supposed. A prey item falling within ränge of the j a w s is immediately grasped. The Euroleon larva apparently never moves from its resting position to actively catch a prey item in the pit. Instead, if the prey is not within ränge, the Euroleon larvae fires sand at the prey item by violently flipping its body upwards so that sand resting on its head and jaws is directed at the prey. Five or six "flicks" in rapid succession are typically deployed and this is usually adequate to dislodge the prey and cause it to fall within ränge. If this fails the larvae will try again with a further series of "flicks" but intersperses these with rest periods of up to 15 seconds. Ant-lions feed on live prey by piercing the prey body with their suctorial mouthparts. After initially grasping the prey item, Euroleon larvae struggle to subdue it by pulling it under the sand. Smaller prey items are usually taken completely below the surface, though larger ones are typically pulled down just sufficient to prevent their struggles. Exceptionally large items, such as cantharid beetles are pulled to the base of the pit and held vertically (either by the head or by the tip of the abdomen) so that the legs have no purchase on the substrate.

Observations on the wild stock Observations of feeding per se are minimal and scarcely significant - all involved ants. However, sieving of the sand in the centre of larger colonies revealed numerous dry exoskeletons of a large number of presumed prey items. These are listed in Table 2, below.

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ANT-LION IN ENGLAND

Taxon ARACHNIDAE ARANEAE Tetragnatha sp. CRUSTACEA ISOPODA Armadillidium vulgare Oniscus asellus Philoscia muscorum Porcellio scaber INSECTA HETEROPTERA MIRIDAE

Code*

Notostira elongata

F

F

D D F D

Taxon COLEOPTERA CANTHARIDAE Cantharis rufa Rhagonycha fulva HYMENOPTERA FORMICIDAE Formica fusca Lasius flavus Lasius niger Myrmica rubra Myrmica ruginodis

75 Code*

F F

F F F F F

Table 2: Natural prey of Suffolk Euroleon nostras * Key: F = larvae seen feeding on prey species D= dessicated exoskeletons sieved in significant numbers from sand within colony Why have ant-lions been overlooked in England? There are a number of reasons why a resident population of large insects such as Euroleon nostras may have been overlooked. Some of these are now summarised: The bulk of the present English population is located in an area of an RSPB nature reserve to which access is restricted. If the meta-population theory is correct, then the outlying "satellite" nesting aggregations are likely to be temporary in nature and may not have been present until quite recently when the Minsmere locus had achieved a critical density. In accessible areas, wandering off the footpath is actively discouraged and, since most of the visitors to the site are specialist bird-watchers, this rule is largely adhered to. Additionally, most visitors to the colony area are either looking upwards for birds or are engaged in site management work and are less likely to notice the pits. In this context, it is probably significant that the pits were first encountered during a specific entomological survey of the site. In its relatively recent history, the RSPB has tended to discourage collecting of insect specimens on its reserves and as a consequence, few entomologists are likely to have visited Minsmere specifically for entomological purposes for fear of being challenged or perhaps even evicted. More recently, however, there has been a more enlightened approach to invertebrate recording on RSPB reserves and, as just mentioned, it was during an entomological survey of the Suffolk Sandlings in 1996 that the larval pits were first encountered at Minsmere. The adult insects are nocturnal. After emergence and drying out, they take to

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the wing and assemble in the upper branches of trees, especially pine trees where mating takes place (Yasseri & Parzefall, 1996). Dßring oviposition, the adult females carefully select suitable sites and test these by tapping the substrate with the tip of the abdomen; it thus follows that most of the night is spent either Walking or engaged in short, probably ground level, flights between potentially suitable areas. Female dispersive flight, which is probably not a regulär feature in any case, may be the only occasion, apart from travel between pine tree and ground, that any "real" flight is engaged in; such flight is likely to explain the 1988 Corton record, though the sex of that specimen is not recorded. It is therefore no surprise that, in general, the adult insects appear to be difficult to attract to mercury vapour lights run by lepidopterists. It is the present author's contention that Euroleon nostras is a permanent resident in England, representing a relic of a population that was formerly more widely distributed over the Suffolk Sandlings, at the extreme north-west limit of its distribution within the Western Palaearctic Region. The English population of Euroleon nostras is centred firmly on the Minsmere RSPB Nature Reserve/Dunwich Heath National Trust site on the Suffolk Sandlings. Thirty-seven of the 42 known colonies fall within this area which can be confined in a rectangle 3-7 kilometres x 1-9 kilometres. The most northerly colony is at Walberswick National Nature Reserve and the most southerly on Blaxhall Common; only 20-5 kilometres separate these two extreme satellite sites. Translated to grid squares, the insect is known from only three tenkilometre squares of the National Grid. It is unlikely that the discovery of larval pits at new sites on the Suffolk Sandlings will significantly alter this Situation. The species has been a resident in this area of England since at least 1986, almost certainly considerably longer, and in view of the thorough searching of the Suffolk Sandlings area during 1997 by several field workers, it is judged rather unlikely that E. nostras has a distribution in this part of East Anglia particularly greater than that here presented. It is probable, however, that at least a few colonies have been overlooked within the main centre of distribution and it is equally likely that more isolated colonies away from the Minsmere/Dunwich Heath focus could be turned up. However, it is perhaps of significance that several apparently suitable sites did not hold larvae during 1997; this is taken as further evidence of a numerically small population. This pattern of larval "colony" distribution fits well with the supposition of Yasseri & Parzefall (1996) that some females may disperse over moderate distances to lay eggs in new areas but that they are not particularly capable of flying excessive distances. The Suffolk population undoubtedly represents a Single meta-population. A restricted availability of suitable egg-laying habitat forces females to lay their eggs in sites which are already occupied by larvae, accounting for the mixed age larval stnicture in these aggregations. Occasional dispersive flights account for the outlying colonies away from the Minsmere/ Dunwich locus. Predation of ovipositing females and of newly emerged adults of both sexes by larvae acts as a natural mechanism maintaining a low population in a small area. It is not considered at all likely, therefore, that the existing meta-population will expand over a particularly wider area and this in

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turn renders the English population vulnerable to human interference of the habitat. It is unlikely, but not considered beyond the bounds of possibility, that quite separate relic populations of this species may persist undetected in other suitable areas of the south-east of England. However, Yasseri & Parzefall (1996) have shown that the adult insect is arboreal and has a particular affinity with pine trees, with large, lone trees being an apparent requirement for mating. If this Situation also applies in England, then it is most unlikely that further populations await discovery away from the Suffolk Sandlings, with the possible exception that some areas of Breckland might be suitable.

Further work required Although this report presents previously unknown Information, the reality of the Situation is that we know remarkably little about this large and distinctive insect which has probably always been a resident in eastern England.

The following work is regarded as essential to understanding the Suffolk ant-lion to aid a Species Recovery Programme. Distribution •

Carry out in-depth surveys of potentially suitable habitat on the Suffolk Sandlings and on the coastal strip area of Norfolk, Suffolk and Essex, including sites examined in 1997 but lacking in larvae as well as sites where earlier sightings were not validated in 1997, to confirm or otherwise the restricted distribution of the species;

Carry out similar surveys in other areas, specifically Breckland to confirm or otherwise the absence of further relic populations. Population level • Monitor all known colonies as far as practical to assess any fluctuations in population and to establish whether such colonies are permanent or whether they may be in different positions from year to year; • Ecology • Correlate any additional discoveries at new localities with the presence or absence of pine trees; • Investigate through field Observation (including pitfall trapping around larval aggregations to assess food availability) and through a captive breeding programme the natural foods of E. nostras in Suffolk and assess these in the context of its conservation; • Assess through field survey the nature and extent of natural predation and other factors which may reduce numbers of E. nostras in Suffolk at all of its developmental stages; •

Assess the habits of adult insects, including an in-depth appraisal of the reasons for failure to detect the presence of these at times when populations are theoretically high;

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Conservation •

Monitor and assess any spread of the insect into any adjacent areas that have been artificially prepared (eg., cleared of Vegetation) with the express purpose of encouraging ant-lions to lay eggs there;

Formulate a broad land-management policy for the conservation of Euroleon nostras on the Suffolk Sandlings;

Acknowledgements Several people have been investigating or taking an interest in the Suffolk ant-lions independent of the present survey. It has been a pleasure to meet with these people and discuss the various aspects of the project with them. I am particularly pleased that all were immediately Willing to share their own observations and results and that it has therefore been possible to draw together in this report, I would therefore like to thank, in alphabetical order: Anglia Television, Steve Clarke (Suffolk Wildlife Trust), Richard Cottle, Nigel Cuming (North-east Essex Coleoptera Recorder), Mike Edwards (Entomological Consultant), Peter Harvey (Essex Hymenoptera Recorder), Penny Hemphill (MSc project Student), Rob Macklin (RSPB North Warren Reserve), Howard Mendel (Ipswich Museum), Simon Moss (National Trust, Dunwich Heath), Geoff Welch (RSPB Minsmere Reserve), Hilary Welch (RSPB Minsmere Reserve), David Wilson (Entomological Photographer), Richard Wilson (RSPB Minsmere Reserve). I also wish to express my thanks to the following people, also presented in alphabetical order, for their help in the project through pointing me to literature references and/or discussing the results with me: Dr Horst Aspöck (Vienna, Austria), Dr Charlie Gibson (Oxford, UK), Dr Andreas Hilliges (Pisa, Italy), Dr Agostino Letardi (Rome, Italy), Dr Roberto Pantaleoni (Sassari, Italy). References Aspöck, H., Aspöck, U. & Holzel, H. (1980). Die Neuropteren Europas. Goeke & Evers, Krefeld. Cottle, R„ Edwards, M. & Roberts, S. (1996). Euroleon nostras (Fourcroy, 1785) (Neur.: Myrmeleontidae) confirmed as breeding in Britain. Entomologist's Ree. J. War. 108: 1-5. Doughty, C. G. (1931). The ant-lion in Suffolk. Trans. Suffolk Nat. Soc.,. 1: 228-229. Killington, F. J. (1932). The ant-lion , Myrmeleon formicarius Linn. (Neur.) in Britain. J. ent. Soc. South of England. 1: 22. Mendel, H. (1996). Euroleon nostras (Fourcroy, 1785) a British species and notes on ant-lions (Neuroptera: Myrmeleontidae) in Britain. Entomologist's Ree. J. War. 108: 1-5. Morley, C. (1931). The ant-lion in Suffolk. Footnote. Trans. Suffolk Nat. Soc. 1: 229. Yasseri, A.M. & Parzefall, J. (1996). Life cycle and reproduetive behaviour of the antlion Euroleon nostras (Geoffroy in Fourcroy, 1785) in northern Germany (Insecta: Neuroptera: Myrmeleontidae). in Canard, M., Aspöck,

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H. & Mansell, M. W. (eds.) Pure and Applied Research in Neuropterology. Proceedings of the fifth international symposium on Neuropterology, Cairo, Egypt, 1994. pp. 269 288. Colin W. Plant Consultant Entomologist 14 West Road Bishops Stortford Hertfordshire CM23 3QP False Truffles One of the first questions to be asked at any "fungus foray" is - "Where can we find truffles ? " It is not an easy question to answer in Suffolk. Arthur Mayfield recorded truffles, both Tuber aestivum and T. ruflim, at Mendlesham in East Suffolk. I have only ever found a partially eaten specimen of T. aestivum in beech woodland on the South Downs, near Stanmer in Sussex. Readers will all know of hunting for truffles (T. melanosporumyill. ) with pigs or dogs in France, and that a good truffle is sometimes claimed to be worth its weight in silver. A whole chapter of John Ramsbotttom's book "Mushrooms and Toadstools" is devoted to truffles and is strongly recommended. T. aestivum is brownish black and covered in pyramidal warts, whereas T. rufum is reddish-brown and only minutely warted. The interior of both species show much branched pale veins. Probably the commonest "nontruffles" found are the earth-balls, Scleroderma spp. ( Heathcote. 1995 ) but root galls produced by the oak apple cynipid, Biorhiza pallida, probably come second. These galls are brown and roughly spherical. The little wasp produces "oak apples" in its unisexual generation. On 2Ist April 1998 I was brought a "non-truffle" which was new to me. Paula Horrabin found it in her garden in Great Barton, near Bury St. Edmunds, amongst soil litter. It proved to be Melanogaster varieqatus var. broomeianus, a Gasteromycete, related to stinkhorns and puffballs, and not an Ascomycete like a truffle. The specimen was roughly rounded, reddish-brown and felty in texture. Inside it was purple-black and divided into irregulär Chambers with white walls. This looked much more like a truffle than any earthball. It had a streng smell which is difficult to describe. I sent a piece of it to Martin Sanford and he told me he could not keep it in his office for long. The only record of this fungus in Martin and Pam Ellis' "Fungi and Slime Moulds in Suffolk" (1988) is from nearby Elmswell, reported by P. W. Glassborrow. References Ellis, M. & P. (1988). Fungi and slime moulds in Suffolk. Suffolk Naturalists' Society, Ipswich. Heathcote, G.D. (1995).Puffballs, earthballs and earthstars. Trans. Suffolk Nat. Soc., 31, 57. Ramsbottom, J. (1953 ). Mushrooms and toadstools. Collins, New Naturalist, London. G. D. Heathcote

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/ i Plate 6: Ant-lion (Euroleon Minsmere, 1997 (p. 71).

nostras

Geoffroy in Fourcroy, 1785) larva at


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