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BR. J. ENT. NAT. HIST., 23: 2010 77 THE AND ASSEMBLAGES (: ACULEATA) OF SOME KEY HEATHLAND SITES IN THE REGION

STEVEN J. FALK Warwickshire Museum, Market Place, Warwick CV34 4SA [email protected]

ABSTRACT A description of the modern bee and wasp assemblages of seven heathlands in the British West Midlands region at the end of the twentieth century is given with an indication of historic losses and recent gains. The characteristics of the various habitats that are typically present at such sites and the way that these are utilised by and are discussed.

INTRODUCTION Despite the ravages of agricultural improvement and the substantial growth of towns and cities within the British West Midlands Region (sensu modern Herefordshire, , Warwickshire, , and the unitary districts such as Birmingham and that formed the former West Midlands County), patches of lowland heathland still remain. However, huge losses have been incurred (90% in Staffordshire over the past 200 years according to DEFRA, 2003) and with the exception of the remaining sites are mostly small and isolated. Today, the Region supports approximately 3000 hectares of ‘heathland’ (DEFRA, loc. cit.) of which nearly 2000 hectares fall within modern Staffordshire, 500 hectares in the former West Midlands County, 108 hectares in modern Worcestershire, and the remainder in Herefordshire, Shropshire and Warwickshire. The precise definition of heathland used by the various sources varies from ericaceous dwarf-shrub communities in the strictest sense to the larger mosaics of heathers, scrub, bracken and grassland that often occur in association. Smaller patches of heather can also be found elsewhere within acid grassland, woodland rides and post-industrial sites, especially in areas with base-poor geology that historically supported heaths and commons (often reflected by place-names containing ‘Heath’ e.g. Short Heath in Birmingham), though these small patches rarely support heathland . West Midlands heaths have attracted entomologists for well over a century, as revealed in the relevant chapters of Victoria County Histories e.g. Martineau (1904) for Warwickshire and Jourdain (1908) for Staffordshire. However, few detailed surveys of bees and wasps at these sites have taken place until the late twentieth century and the fauna has traditionally been much less understood than that of southern counties such as Dorset, Hampshire and Surrey, or that of East Anglia. Since 1990 the author and a number of workers including Dr Michael Archer, Jon Webb and Andy Jukes have made repeat visits to various sites of known or potential value and a number of papers and technical reports have already been produced (Falk et al., 1996; Falk & Lane 1999; Archer, 2002, 2004, 2006: Falk & Webb, 2002; Jukes, 2004). This paper collates these published and unpublished data and attempts to provide a better understanding of the contribution that heathland in this part of Britain makes to our bee and wasp fauna. 78 BR. J. ENT. NAT. HIST., 23: 2010

METHODS

Survey sites Seven sites were surveyed: (i) Baddesley Common, north Warwickshire (SP274978) (Fig. 1) – a tiny heathland of a few hectares, studied by the author on several dates in 1995. (ii) Barlaston/Rough Close Common, north Staffordshire (SJ925395) – a medium- sized (approximately 21 ha) mosaic of dry heathland, wet heathland, bracken and woodland containing a small sand quarry. Studied by the author on several dates in 1994 and 1995. (iii) Devil’s Spittleful SSSI, Worcestershire (SE8270) – a 60 ha site with much sandy heathland, studied by Dr Archer on numerous dates between 1990 and 1997. (iv) SSSI, mid Staffordshire (SK050113) (Fig. 2) – an 80 ha site near Chase Terrace supporting both dry and wet heathland, and also containing a sandstone quarry. Studied by the author on four dates in 1994/95, with subsequent visits by Dr Archer and A. Jukes up to 2006. (v) Hartlebury Common SSSI, Worcestershire (SO823706) – an 87.5 ha site with much dry sandy heathland, studied by Dr Archer between 1990 and 1997. (vi) Highgate Common Park, (SO838899) (Fig. 3) – a 94 ha mosaic of dry heathland, acid grassland and woodland. Studied by the author, Dr Archer, A. Jukes and J. Webb on numerous dates between 1996 and 2006. (vii) Sutton Park National Nature Reserve, north Birmingham (SP1096) (Fig. 4) – a 989 ha mosaic of dry heathland, acid grassland, wet heathland and mire, carr and woodland. Studied by the author on numerous dates in 1996 and 1997.

Survey methodology The nature of the surveys varied between the recorders. Much of it took the form of inventories that attempted to produce good species lists. Some visits, such as those of Dr Archer, concentrated purely on bees and wasps, whilst many of the author’s visits also covered Diptera, Odonata and butterflies, which influenced they way in which bees and wasps were sampled (with a heavy emphasis on sweeping). The author’s inventory surveys were also structured to produce separate species lists for various vegetation zones and microhabitats within the study sites e.g. bare sand, heather stands, acid grassland, wet heath, bracken, gorse stands and flowery, shrubby heathland ‘verge’. By contrast, work carried out by the author at Highgate Common between 2000 and 2002 was mostly autecological in nature, concentrating on observing the way in which the scarcer species were using the site. In all surveys, special attention was paid to both nesting and foraging areas. Examination of sandy footpaths and any sand faces across a range of dates revealed many of the fossorial species present and provided an indication of population strengths. Key forage plants were checked at the peak of their flowering, including heathers Calluna vulgaris and Erica species, bilberry Vaccinium myrtillus, cat’s-ear Hypochaeris radicata, mouse-ear hawkweed Pilosella officinarum, grey Salix cinerea, wild cherry Prunus avium, blackthorn , hawthorn , umbellifers such as hogweed sphondylium, brambles, thistles Carduus and Cirsium species, common knapweed Centaurea nigra, common ragwort Senecio jacobaea, common ’s-foot trefoil , vetches Vicia species and harebell Campanula rotundifolia. For genera such as , Lasioglossum, BR. J. ENT. NAT. HIST., 23: 2010 79

Fig. 1. Baddesley Common, Warwickshire, a small areas of heathland with a limited fauna of bees and wasps.

Fig. 2. Gentleshaw Common, north Staffordshire – one of several sites with an old sandstone quarry which is much exploited by fossorial species. 80 BR. J. ENT. NAT. HIST., 23: 2010

Fig. 3. Highgate Common, south Staffordshire – the most exceptional of the sites with a list of 130 bees and wasps.

Fig. 4. Sutton Park, north Birmingham – a large suburban heathland with wet and dry heath, of moderate value for bees and wasps. BR. J. ENT. NAT. HIST., 23: 2010 81 , Diodontus and smaller care was taken to obtain reasonable samples, as several species that are indistinguishable in the field can be present. Prolonged sweeping using a long-handled net was employed by the author and proved to be one of the most efficient sampling methods over mixed nesting colonies and flower-rich habitats. It was also particularly effective for obtaining pompilids, small crabronids such as Harpactus tumidus (Panzer) and dimidiatus Jurine and also the tiphiid Tiphia minuta Vander Linden in short or sparse vegetation, as these can be time-consuming insects to capture individually and are not always easily spotted in the field. Visits by the author (also covering Diptera) generally lasted for 5–6 hours and every effort was made to exploit reasonably warm and sunny weather as overcast or wet conditions severely affect the activity of bees and wasps. Visits by Dr Archer lasted from 2–3 hours and involved use of a short- handled kite net. The data have been analysed in a number of ways. The presence of Red Data Book, Nationally Scarce and Regionally Scarce species has been used to obtain a standard rarity score for each site by assigning points to the various rarity 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 gradings for Red Data Book and Nationally Scarce species were taken from Falk (1991) but some have been adjusted to allow for the fact that some gradings are now known to be strongly misleading, either due to species increasing their range and frequency in recent decades, or because they were under-recorded in the past (within the Appendix an asterix has been placed against the obviously misgraded species and a bracketed regrade suggestion that is more realistic given afterwards). Rarity scores quoted in Table 1 are based on these adjusted grades. Regional scarcity (sensu West Midlands) was assessed using the national atlases published by the Bee, Wasp and Ant Recording Society (e.g. Edwards 1997 & 1998, Edwards & Telfer, 2001 & 2002), also personal data including the official dataset for Warwickshire bees and wasps, the on-line Staffordshire checklist (available at www.staffs-ecology.org.uk). Dr Archer has also worked out a species quality score (here termed the Archer species quality score) that he has developed using the distributions and frequencies of different species in the same national atlases (see Archer, 1995 for methodology). An effort was also made to assess the quality and characteristics of the bee and wasp assemblages at these sites by examining the presence of key indicator species. The categories used were: . Species with a strong or obligate requirement for heathers (marked ‘Er’ in the Appendix) e.g. Andrena argentata Smith, A. fuscipes (Kirby), (L.) and the cleptoparasite Epeolus cruciger (Panzer), the last because it seems to be entirely parasitic upon C. succinctus in the West Midlands. . Other species strongly associated with heathland or acid grassland in the West Midlands i.e. ‘heathland-loving’ even if they do not specifically require heathers (marked as ‘HL’ in the Appendix). Not all of these species are so strongly associated with heathland and acid grassland in other parts of Britain (Andrena humilis Imhoff for example regularly occurs in chalk grassland on the South Downs). . Coastal-biased species, a special sub-group of the last category for four species that typically occur on dunes, but can colonise inland on sites with particularly loose sands, namely Dasypoda hirtipes (Fabr.), Megachile maritima (Kirby), Oxybelus argentatus Curtis and Podalonia affinis (Kirby). Their presence inland can be considered a special attribute of a site in the same way as a comparable plant population might be viewed. 82 BR. J. ENT. NAT. HIST., 23: 2010 Table 1. The number of scarcer species of bee and wasp found at each of the seven sites, rarity scores and Archer species quality scores

Barlaston/ Sutton Baddesley Rough Devil’s Gentleshaw Hartlebury Highgate Park Common Close Spittleful Common Common Common NNR

Total no. of species 57 52 122 112 100 130 92 Red Data Book species 00 01 150 Nationally Scarce species 2215812122 Regionally Scarce species 8 2 31 18 32 29 18 Standard rarity score 260 140 1370 860 1340 1680 460 Archer species quality score 2.1 1.4 2.5 2.3 2.9 3.0 2.1

RESULTS Two hundred and sixteen species of bee and wasp (excluding Apis mellifera L.) were recorded with certainty from the seven sites in the West Midlands and are listed in the Appendix (see Plate 5, Figs 4–6 for notable examples). This is not considered a definitive list, and the recording of certain cleptoparasitic bees and wasps (e.g. Sphecodes rubicundus Hagens and Nysson dimidiatus) at sites without finding their hosts indicated that a level of under-recording still remained at the end of the surveys. The richest site proved to be Highgate Common with 130 species (Table 1). This is currently the longest bee and wasp list for any West Midlands site. Baddesley Common and Barleston/Rough Close Common were the poorest by some margin. The totals for all sites are given in Table 1. Many rare and scarce species were encountered. Adjusted for misgrading, this included six reasonably valid Red Data Book species, 26 Nationally Scarce species and 48 Regionally Scarce species. These species are highlighted in the appendix and the total number recorded at each site, plus the rarity score derived from these is shown in Table 1. Species new to the Staffordshire list included Diodontus insidiosus Spooner, Dolichovespula saxonica (Fabr.), Lasioglossum brevicorne (Schenck), Nysson dimidiatus, N. trimaculatus (Rossius), Sphecodes niger Hagens, S. reticulatus Thomson and S. rubicundus. Elampus panzeri (Fabr.) was new to the Warwickshire list. signata Jurine had not been recorded in the region for many years. The records for L. brevicorne and D. insidiosus were substantial extensions to the known ranges in Britain. In terms of heathland habitat indicator species, Highgate Common, Hartlebury Common and Devil’s Spittleful proved to be much richer for these than the others (Table 2).

Table 2. The number of heathland habitat indicator species at each of the seven sites

Barlaston/ Sutton Baddesley Rough Devil’s Gentleshaw Hartlebury Highgate Park Common Close Spittleful Common Common Common NNR

Species requiring heathers 30 53 432 Other heathland lovers 3 4 23 16 26 32 13 Coastal-biased species 00 30 400 Total indicators 6 4 31 19 34 35 15 BR. J. ENT. NAT. HIST., 23: 2010 83

DISCUSSION Factors affecting comparative richness With the exception of Hartlebury Common and Devil’s Spittleful, which were recorded exclusively by Dr Archer in a comparable manner, all the other sites had experienced different levels of recording, which made comparison rather difficult based on gross data. However, comparisons become easier if data are viewed more selectively. The initial survey of Barlaston/Rough Close Common, Gentleshaw Common and Highgate Common by the author in 1994 and 1995 was done in a comparative manner involving four 5–6 hour visits to each site in fine weather on similar dates. It revealed that Highgate was relatively rich (77 species), Barlaston/ Rough Close Common was relatively poor (52 species) and Gentleshaw was intermediate (64 species). The total recording effort for Highgate Common (which extended beyond 1995) and Sutton Park was also broadly similar (about a dozen visits on varied dates). It was clear that Highgate was by far the richer of the two, and had good populations of many key species, including the relatively conspicuous Bombus jonellus (Kirby), Epeolus cruciger (Panzer) and sabulosa (L.) that could not be detected at Sutton Park despite targeted searches. Another way sites were compared was by examining data from single dates during key flowering peaks e.g. August when the heather is fully in flower, or April when vernal species are most active. For sites visited by the author, this supported a ranking of least rich to most rich sites in the order of Baddesley Common, Barleston/Rough Close Common, Sutton Park, Gentleshaw Common and Highate Common. Careful examination of the data for Devil’s Spittleful and Hartlebury Common (Archer, 2002, 2004) suggested that they both approach Highgate Common in richness, but require further spring visits to sample Andrena and Nomada species in detail as these seemed to be rather poorly represented in the site lists. A number of factors are likely to be influencing the richness of a site: . Site size and the provision of key habitats. Larger sites with more plentiful open ericaceous heathland alongside a variety of other habitats (especially reasonable quantities of flowery ‘verge’) seemed to be richer than smaller sites with only a limited provision of open heathland (e.g. Barlaston/Rough Close Common and Baddesley Common) or limited flowery verge. But size was clearly not the only factor at play, because the very large Sutton Park (989 ha) was clearly not as rich as the moderately-sized Highgate Common (94ha). . Geographical location and altitude. Gentleshaw Common and Barleton/Rough Close Common are elevated sites ecologically akin to the nearby Cannock Chase, with upland heathland elements in their vegetation. Highgate Common, Hartlebury Common and Devil’s Spittleful which are located much further south and at lower altitude, have a character much closer to typical southern English lowland heathland and supported a much larger number of southern-biased species. Sutton Park seemed to fall between the two categories. Within the West Midlands, there seems to be a tailing off of aculeate diversity in heathland as one moves north and on to higher land. . Geology, topography and ground conditions. Geology and hydrology influences the friability of sandy ground and its suitability for fossorial species, some of which appear to be very selective about where they nest. Topography can influence the presence of warm, south-facing slopes and banks that are favoured by many species. At all the sites studied by the author, there were clear concentrations of mixed-species nesting colonies along certain stretches of footpath or in old 84 BR. J. ENT. NAT. HIST., 23: 2010 quarries. Here sandy ground was dry, relatively firm and fully exposed to the sun, but very loose sand, damper or semi-shaded stretches of sandy footpaths had much lower interest. . Site history. Highgate Common had experienced a long history of motorcycle activity prior to becoming a country park. Whilst this might have been viewed as incompatible with nature conservation at the time, it has left a legacy of much sandy ground that clearly suits many ground-nesting species. At Gentleshaw Common, by contrast, the relatively pristine heathland, much of which has a wet or humid nature, has considerably less good nesting habitat for bees and wasps. A history of quarrying at Gentleshaw Common, Barleston/Rough Close and Baddesley Common provides important nesting habitat and is likely to be responsible for stronger populations of many species and greater overall richness than might otherwise exist. The historical presence of unstable blown sands at Hartlebury and possibly Devil’s Spittleful seems to be the factor underlying the presence of several coastal-biased species there.

Factors affecting the presence of rare species and fine habitat indicator assemblages Table 1 clearly illustrates the considerable importance of Highgate Common, Hartlebury Common and Devil’s Spittleful for scarcer bees and wasps, many of which were unknown from any other sites in the West Midlands at the time of the surveys e.g. Ammophila pubescens Curtis (Plate 5, Fig. 5), Andrena argentata, A. bimaculata (Kirby), A. nigrospina Thomson (Plate 5, Fig. 6), A. ovatula (Kirby), Dasypoda hirtipes, Diodontus insidiosus, Hedychridium cupreum (Dahlbom), Lasioglossum brevicorne, L. parvulum (Schenck), Megachile maritima, haemorrhoidalis (Fabr.), Nomada fulvicornis Fabr., Oxybelus argentatus Curtis, Podalonia affinis (Kirby), gracilis Haupt and Sphecodes reticulatus Thomson. Geographical location, coupled with reasonably large site size and the presence of particularly fine sandy nesting habitat seemed to be important factors underpinning this. These factors are also likely to be responsible for the relatively large number of heathland indicator species, and it appears that relatively small and isolated fragments of heathland such as Barleston/Rough Close and Baddesley Common had much less capacity for supporting typical heathland species than larger sites. Andrena humilis, Lasioglossum fratellum (Pe´ rez) and Priocnemis schioedtei Haupt are notable for being capable of clinging on to small or degraded fragments of heathland or acid grassland lacking heathers. Of the rare species encountered, two deserve particular mention. The colony of Andrena nigrospina at Highgate is one of only a handful currently known to exist in Britain (Fig. 5). The autecological work by the author reaffirmed that it is a univoltine species peaking in high summer in contrast to the almost identical but bivoltine A. pilipes (Fabr.) that predominates in coastal districts of southern Britain. The autecological study also revealed that the univoltine population of Nomada fulvicornis attacking A. nigrospina is consistently different in appearance to the bivoltine population of N. fulvicornis attacking the closely-related Andrena bimaculata at the same site (see Falk, 2004). DNA analysis is required to more fully understand the taxonomic relationship between the two forms. More recent work by Andy Jukes and others at a colony in Worcestershire has revealed that A. nigrospina may rely heavily on wild radish Raphanus raphanistrum at arable field margins for pollen (Jukes, 2009). BR. J. ENT. NAT. HIST., 23: 2010 85 The Rarity Score approach versus the Archer Species Quality Score The ranking of sites based on scarcer species comes out the same using both approaches, but a poorer sensitivity of the Archer Species Quality Score is revealed in the identical scores obtained for Sutton Park (92 species and a Rarity Score of 460) and Baddesley Common (57 species and a Rarity Score of 260) and also for the relative scores as a whole. It would be difficult to conclude that Sutton Park and Baddesley Common are similar in quality using raw data, and this perhaps highlights the dangers in converting raw data to scores and indexes using equations. This suggests that it is better to strive towards comparable levels of recording at different sites that need to be compared or to focus on comparable elements within existing data for those sites.

THE VALUE OF INDIVIDUAL HEATHLAND HABITATS FOR BEES AND WASPS There was considerable variation in the importance of the different habitats constituting heathland sensu lato for bees and wasps, and this was greatly influenced by the time of year and the precise floristic composition of the various vegetation types. The numbers of species recorded from the various habitats surveyed by the author during his studies are shown in Table 3.

Fig. 5. A mining bee nest (suspected Andrena nigrospina) at a path edge. 86 BR. J. ENT. NAT. HIST., 23: 2010 Table 3. The number of species recorded in the various habitats present at Barlaston/Rough Close, Gentleshaw, Highgate and Sutton Park, based on data from the 1994–95 Staffordshire survey and the 1996–97 Sutton Park survey.

Barlaston Sutton Rough Gentleshaw Highgate Park Close Common Common NNR

Heather-dominated areas 5 19 26 19 Dry, acid grassland 10 5 28 35 Wet and humid heath 6 9 – 18 Bare or sparsely-vegetated sand 36 48 50 31 Bracken stands 4 6 10 7 Heath verge (Sutton Park not covered) 33 47 52 – Gorse and broom scrub (Gentleshaw not covered) 2 – 11 11 Pool, mire, water edge and marshy grassland 2 0 1 27 Coniferous woodland (Sutton Park only) –––3 Dry, broad-leaved woodland (Sutton Park only) – – – 32 Wet woodland & carr (Sutton Park only) – – – 13

Dashes signify a lack of data.

Heather-dominated areas For much of the year, heather-dominated vegetation was a fairly unproductive habitat, but during the flowering of ling Calluna vulgaris (typically August and September), bell heather Erica cinerea (mid June to September) and cross-leaved heath Erica tetralix. (July to September), such areas were heavily exploited by foraging insects of many sorts. These included two oligolectic bees, Colletes succinctus and Andrena fuscipes, which forage mainly on heathers; also large numbers of Bombus, Lasioglossum, Sphecodes and Nomada rufipes Fabr. Predatory wasps also used heather flowers as a bountiful source of prey, with rybyensis and Philanthus triangulum (Fabr.) (Plate 5, Fig. 4) hunting for bees whilst Mellinus hunted flies. The pompilid Priocnemis schioedtei was not observed visiting flowers but seemed to favour a heather-acid grassland/bare sand mosaic and may have been hunting spiders that specifically require this mosaic. also favoured ericaceous vegetation at Highgate Common and presumably finds its caterpillar prey here or in the grassy interstices. But its requirement for heather is not obligatory in contrast to the much rarer A. pubescens which was found at Hartlebury (Plate 5, Fig. 5). Heather-dominated vegetation also supported some of the other useful forage flowers listed under dry acid grassland below, plus (at Highgate) plentiful western gorse Ulex gallii.

Bare sandy ground Bare or sparsely-vegetated sandy ground is an essential feature for the many fossorial bees and wasps plus the large number of cleptoparasites they support. This includes other bees and wasps, flies such as Bombylius, Metopia, Macronychia, Senotainia and Leucophora, plus an important population of the oil beetle Meloe proscarabaeus L. at Highgate. As stated above, bare ground was observed to vary significantly in character and this greatly influenced the distribution of nesting by a species and the richness of a mixed nesting colony. This was particularly evident at Highgate, where two relatively small sandy areas supported particularly rich nesting BR. J. ENT. NAT. HIST., 23: 2010 87 colonies including the main nesting areas for many of the scarcer species present (e.g. Andrena nigrospina, A. bimaculata and Philanthus triangulum), even though a much larger network of sandy paths was available. Factors that appeared to influence the distribution of nests included dampness of the ground, its precise composition (e.g. the balance of sand versus clay or humus/peat and particle size), the pattern of trampling, degree of shading, width of a path, aspect, exposure to prevailing winds and probably the age of a path. Most fossorial species favour ground that is fully exposed to the sun, faces south, is easily accessed from foraging or hunting areas, and is not excessively churned up by bikes or horses. It should be noted that within footpaths and other well-trampled areas, a gradient of disturbance will typically occur between bare sand and adjacent ericaceous vegetation or acid grassland, and many species favour intermediate stages within this gradient, including firm bare ground, mossy areas or short turf. A few species were found to tolerate more shaded areas e.g. Andrena fucata Smith and A. scotica Perkins (often betrayed by their Nomada parasites) and Haupt. The less trampled margins of footpaths can also be a valuable source of flowers such as dandelions and daisy Bellis perennis, which are visited by certain Andrena, Lasioglossum and Nomada species. As noted earlier, bare ground comprising sand that was once mobile blown sands (notably at Hartlebury Common) supported a number of characteristic species that avoid more compacted or finer sands, notably the four ‘Coastal’ species listed in Table 2. Several further species, including Colletes fodiens Geoffroy and Dryudella pinguis (Dahlbom), also tolerate loose sands, but are rather more frequent inland.

Dry acid grassland The value of dry acid grassland for bees and wasps was heavily dependent on the flowers within it and the presence of bare ground. Where floristically poor and dominated by a dense turf of grasses such as wavy hair-grass Deschampsia flexuosa,it remained an unproductive habitat for much of the year. Important forage plants that improved its value included cat’s-ear and mouse-ear hawkweed (especially for Andrena humilis and Lasioglossum species), ragwort (many bee and wasp genera), common bird’s-foot-trefoil (especially for Bombus, Megachile, Hoplitis claviventris (Thomson) and certain Andrena species), harebell (for Melitta haemorrhoidalis), bilberry (for Andrena lapponica Zetterstedt and certain Bombus species), tormentil Potentilla erecta (for Andrena tarsata Nylander, Lasioglossum species and smaller Andrena), heath bedstraw Galium saxatile (for Lasioglossum species and smaller Andrena), rosebay willowherb Chamerion angustifolium (especially for Bombus, Megachile, Nomada and Philanthus), patches of heathers, gorses, broom and any bushes of rowan, hawthorns or Prunus species.

Bracken stands Dense bracken Pteridium aquilinum was one of the least productive parts of the sites, supporting few flowers before the bracken fronds had developed and holding very little interest for bees and wasps once they had fully formed. In spring, male Andrena and Priocnemis species were sometimes observed using sunlit dead bracken litter for swarming, presumably because it produces warm microclimates.

Wet and humid heath, mire and water edge Again, the value of these moist habitats depended on the flowers present and they were least productive for bees and wasps when dominated by dense tussocks of 88 BR. J. ENT. NAT. HIST., 23: 2010 purple moor grass Molinia caerulea. Valuable flowers noted were cross-leaved heath, devil’s-bit scabious , Dactylorhiza orchids and at Sutton Park, marsh thistle Cirsium palustre, meadow thistle Cirsium dissectum, wild angelica , water mint Mentha aquatica, great willowherb Epilobium hirsutum and also some of the plants listed under dry acid grassland. Bumblebees were especially numerous in flowery wet heath.

Flowery ‘verge’ habitat This habitat was typically the most floristically diverse part of a heathland and tended to coincide with areas of enriched soil around the edges of sites and along the verges of intersecting roads or tracks. Valuable forage plants present in verge habitat include umbellifers (e.g. common hogweed, cow parsley Anthriscus sylvestris and hedge parsley Torilis japonica), thistles, black knapweed, dandelions, vetches, clovers Trifolium spp., dead nettles Lamium spp., comfreys Symphytum spp., buttercups Ranunculus spp., ground-ivy Glechoma hederacea. (and any other lamiates), and shrubs such as grey willow, hawthorns, roses, wild cherry, rowan, crab apple Malus sylvestris and brambles. Floristically-rich verge habitat seems to substantially increase the diversity of bees and wasps at a site and the number of scarcer species present. It is also an important source of prey for certain sphecid wasps e.g. Ectemnius species which like to hunt on umbellifer flowers. The paucity of flowery verge habitat at Sutton Park was regarded as one of the reasons for the relatively poor diversity of bees and wasps recorded there.

Gorses and broom The flowers of these shrubs attracted certain Bombus, Andrena, Lasioglossum and Halictus species, often in very large numbers. Andrena bimaculata, A. dorsata and A. ovatula seemed particularly dependent upon common gorse in spring and males formed large swarms around flowering bushes. Certain Andrena species regularly swarmed around blossoming broom, most notably the males of the rare A. nigrospina at Highgate, though the females did not appear to forage on it. Females of A. synadelpha clearly were attracted by broom flowers at Highgate and bumblebees visited it too for pollen, though it does not produce nectar. At Highgate, western gorse flowered at the same time as heathers and supported some bumblebee foraging though most bees and wasps preferentially visited heathers.

Woodlands, scrub and carr Relatively little bee and wasp activity was observed to take place within the interior of shaded woodland, dense scrub or carr, though these habitats benefited the fauna in a number of ways. The blossoms of , hawthorn, Prunus, Sorbus and Acer species are vitally important for spring flying Andrena species and queen bumblebees. A further valuable feature of woodland, scrub and carr is the dead wood it produces, which provides nesting opportunites for wasps such as Crossocerus, Ectemnius and Pemphredon, plus Osmia and Megachile bees. Sutton Park was the most important in this respect, having a good number of veteran trees, plus some large log piles resulting from regular tree work. Even piles of old building timbers or old wooden sheds (as present in the ranger’s yard at Highgate) supported a variety of dead wood-nesting species. It is important to bear in mind that many bee and wasp species will require several of the above-mentioned habitats to complete a life cycle. For bees with two BR. J. ENT. NAT. HIST., 23: 2010 89 generations, e.g. Andrena bimaculata, the floristic requirements are likely to vary substantially between the spring generation (peaking in April) and the summer generation (peaking in July). If just one of the requirements of one of the generations is lost, the bee could become locally extinct.

EVIDENCE FOR LOCAL LOSSES AND GAINS ON WEST MIDLANDS HEATHLAND It is clear from data gathered by the Bee, Wasp and Ant Recording Society that the bee and wasp faunas of most parts of Britain are changing (e.g. Edwards 1997 & 1998, Edwards & Telfer, 2001 & 2002), and this is affecting many habitats including heathland. These changes involve both loss of species (usually in response to habitat loss and fragmentation) plus gains as certain southern-biased species and recent British colonists expand their ranges north. Most of the sites covered here lack good historic data, but a review of old literature records for insects at Sutton Park (Falk & Lane, 1999) added 21 species to the site list and strongly suggests several species are likely to have been lost including Andrena tarsata Nylander, Bombus distinguendus Morawitz and B. jonellus. Species that appear to have disappeared completely from West Midlands heathland on the basis of published national data include Bombus distinguendus, B. sylvarum (L.), longicornis (L.) and Nomada roberjeotiana Panzer. This has been counter-balanced by the arrival of Philanthus triangulum, Cerceris rybyensis, Dolichovespula saxonica (Fabr.), Lasioglossum brevicorne, Sphecodes niger Hagens and S. reticulatus.

CONCLUSIONS The remaining heathland of the West Midlands contributes substantially to the biodiversity of bees and wasps in Britain, extending the ranges of many species and supporting some critically important individual populations. There is still a great deal to learn about these sites and informed management and monitoring will be crucial for sustaining or improving the quality of the fauna. One of the most positive outcomes of the above studies has been the notification of Highgate Common as a Site of Special Scientific Interest, and also the funding that has been made available to study the autecology of Andrena nigrospina at Highgate Common and a Worcestershire site. Staffordshire Biodiversity Partnership also managed to pursuade JCB to act as the champion for Solitary Bees and Wasps in Staffordshire, and provide some funding for surveys and site management work (see www.sbap.org.uk/ saps/bees and www.sbap.org.uk/haps/lowheath).

ACKNOWLEDGEMENTS The author is particularly indebted to Sue Sheppard (formerly of the Staffordshire and West Midlands Heathland Partnership) for setting up the various surveys led by the author and providing constant support throughout, and English Nature (now Natural ), who financed these surveys. I would also like to thank Dr Michael Archer for permitting use of his data, Roger Hill (former County Ecologist, Staffordshire County Council), Andy Jukes (formerly Staffordshire Wildlife Trust), and my co-surveyors Steve Lane, Jon Webb, Craig Slawson and Mike Bloxham. Maximillian Schwarz (Austria) provided advice on the two forms of Nomada fulvicornis encountered at Highgate Common, and the Natural History Museum, London and Oxford University Museum kindly permitted examination of N. fulvicornis material in their aculeate collections. Thanks are also due to Stefan Bodnar and the Rangers at Sutton Park (Birmingham City Council), the Rangers at 90 BR. J. ENT. NAT. HIST., 23: 2010 Highgate Common, Peter Coxhead (Sutton Natural History Society), the late Pam Copson (formerly Warwickshire Museum) and the staff at Ecorecord (The Biological Records Centre for Birmingham and the Black Country).

REFERENCES Archer, M. E. 1995. Aculeate wasps and bees (Hymenoptera: Aculeata) of Blaxton Common in Watsonian Yorkshire with the introduction of a new national quality scoring system. Naturalist 120: 21–29. Archer, M. E. 2002. The wasps and bees (Hymenoptera: Aculeata) of Hartlebury Common Local Nature Reserve in Watsonian Worcestershire. Proceedings of the Birmingham Natural History Society 28: 2–19. Archer, M. E. 2004. The wasps and bees (Hym., Aculeata) of Devils Spittleful Nature Reserve in Watsonian Worcestershire. Entomologist’s Monthly Magazine 140: 39–49. Archer, M. R. 2006. The wasps and bees (Hym., Aculeata) of the open sandy habitats of Highgate Common in Watsonian Staffordshire. Entomologist’s Monthly Magazine 142: 207–218. Ball, S. G. 1986. Terrestrial and Freshwater Invertebrates with Red Data Book, Notable or Habitat Indicator Status. Invertebrate Site Register 66 (Contract Reports, 637). Nature Conservancy Council, Peterborough. DEFRA 2003. England Rural Development Plan – West Midlands, DEFRA. 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. 2004. The form of Nomada fulvicornis F. (Hymenoptera ) associated with the mining bee Andrena nigrospina Thomson. British Journal of Entomology and Natural History 17: 229–235. Falk, S. & Lane, S. 1999. A survey of the insects of Sutton Park, Birmingham. Falk Entomology unpublished report for English Nature and the Staffordshire and West Midlands Heathland Partnership. Falk, S. & Webb, J. 2002. A survey of the scarcer insects of Highgate Common, Staffordshire (2000–2002). Falk Entomology unpublished report for English Nature and the Stafford- shire and West Midlands Heathland Partnership. Falk, S., Lane, S., Slawson, C. & Bloxham, M. 1996. A Comparative Study of the Invertebrate Assemblages of Three Staffordshire Heathland Sites. Coventry Museum Ecology Unit unpublished report for English Nature and the Staffordshire and West Midlands Heathland Partnership. Jourdain, F. C. R. 1908. Hymenoptera. In: The Victoria County History of Staffordshire 1: 82–86. Jukes, A. 2004. An evaluation of the invertebrate fauna of Highgate Common in relation to other sites within Staffordshire and the West Midlands (Extended version). Report by Staffordhire Wildlife Trust for English Nature. Jukes, A. 2009. Andrena nigrospina – Ecology and conservation of a Red Data Book solitary bee. Report for thee Staffordshire Wildlife Trust. Martineau, A. H. 1904. Hymenoptera. In: The Victoria County History of Warwickshire 1: 73–76. BR. J. ENT. NAT. HIST., 23: 2010 91 + + Park Sutton (continued) + ++ Highgate Common + ++ Common Hartlebury Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley Coquebert) R, HL + + + + + in ) associated species, HL – other species strongly associated with heathland in the Midlands, Coastal – species with a strong (Fabr.) (Magretti) + (Dahlbom) N, HL + + (Spinola) R + + (Latreille (Dahlbom) R, HL Erica Latreille N, HL + + (Wesmael) + + + + (L.) N* (no status) (Scopoli) + + (Schiødte) + + + + Schenck + + (L.) + + (L.) R, HL and (Fabr.) R, HL (L.) R + + Panzer + + + + Vander Linden N* (no status) Calluna Cleptes semiauratus Sapyga quinquepunctata Chrysididae Chrysis angustula Elampus panzeri Mutillidae Myrmosa atra Pompilidae Agenioideus cinctellus Anoplius nigerrimus Hedychridium cupreum Trichrysis cyanea Tiphiiidae Methocha articulata Tiphia minuta Hedychridium ardens anceps Arachnospila trivialis Arachnospila spissa Dipogon subintermedius Episyron rufipes coastal bias. An asterix after the nameACULEATE indicates HYMENOPTERA a misleading national grading. A bracketed grading indicates a more realistic grading. APPENDIX. A fullspecies). list Key: of RDB1, the 2heather bees & ( and 3 wasps – for Nationally Threatened the categories, seven N sites, – highlighting Nationally the Scarce, scarce R – species Regionally and Scarce habitat in indicator the species West (Quality Midlands Indicator Region, Er – 92 BR. J. ENT. NAT. HIST., 23: 2010 + Park Sutton (continued) + Highgate Common +++ Common Hartlebury ++ Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley ller) ¨ (Fabr.) + + + (Mu (Curtis) + + (Scopoli) + + + + + + (Fabr.) RDBK* (no status) + + (L.) (L.) + + + + + (Harris) + + + (L.) Curtis R* (N), Er + + (Retzius) + Haupt N, HL + + + (Shuckard) R + + + + (L.) R, HL + + + Haupt R, HL + + Dahlbom + + + + Haupt N, HL (Fabr.) + + + + + + + (L.) R, HL + + + + + (L.) R, HL + + + (L.) R, HL + + + (Kirby) RDB3* (N), Coastal + (L.) + + + + + + + (continued) L. + (L.) + + + + + + + Priocnemis gracilis Priocnemis parvula Priocnemis schioedtei Priocnemis susterai Vespidae Ancistrocerus nigricornis Ancistrocerus parietum Ancistrocerus trifasciatus Cerceris rybyensis Cerceris arenaria cribrarius Dolichovespula media Dolichovespula norwegica Dolichovespula saxonica Dolichovespula sylvestris Symmorphus bifasciatus Vespa crabro Vespula germanica Vespula rufa Ammophila pubescens Ammophila sabulosa Podalonia affinis Vespula vulgaris APPENDIX. ACULEATE HYMENOPTERA BR. J. ENT. NAT. HIST., 23: 2010 93 + + Park Sutton (continued) ++ + ++ ++ + + Highgate Common + Common Hartlebury ++++ Common Gentleshaw ++ ++ Devil’s Spittleful Common Barlaston/ Rough Close ++++ ++ Common Baddesley ) ´ ´ ´ ´ (Fabr.) + + + + + (Rossi) + + + (Morawitz) N + (Vander Linden) + (Vander Linden) (Fabr.) (Lepeletier & Brulle (Vander Linden) R, HL + + (Olivier) (Wesmael) (Shuckard) + + + Spooner RDB3, HL (Fabr.) (Zetterstedt) N + Lepeletier & Brulle Lepeletier & Brulle (Thomson) + + Fabr. + + + + + + (Panzer) R + + + ? (Fabr.) R + + Lepeletier & Brulle Jurine N* (R) (Dahlbom) R, HL (Vander Linden) R* (N), HL + (Fabr.) R, HL + + + + + (L.) + + + + + + + (Vander Linden) R, HL + + + (Shreber) R, HL + + + (Fabr.) R, HL + + + + (continued) Diodontus minutus Diodontus tristis Dryudella pinguis Nysson dimidiatus Harpactus tumidus Lindenius albilabris Lindenius panzeri Mellinus arvensis Mimesa equestris Mimesa lutaria Mimumesa dahlbomi Crossocerus quadrimaculatus Crossocerus tarsatus Crossocerus wesmaeli Diodontus insidiosus Ectemnius cavifrons Ectemnius cephalotes Ectemnius continuus Ectemnius ruficornis quadrifaciatus Crossocerus ovalis Crossocerus podagricus Crossocerus pusillus Crabro peltarius Crossocerus distinguendus Crossocerus elongatulus Crossocerus megacephalus Crossocerus nigritus Crossocerus annulipes APPENDIX. ACULEATE HYMENOPTERA 94 BR. J. ENT. NAT. HIST., 23: 2010 + + + + Park Sutton (continued) + Highgate Common +++ + Common Hartlebury + +++ ++ + Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley (Panzer) R, HL + + + + + Dahlbom R, HL Smith + sensu lato Lepeletier & Serville + (Fabr.) RDB2* (R)(Scopoli) + + + + + + Say (Schuckard) (Kirby) + + + (Fabr.) + + + + Schuckard + + Curtis N, Coastal + + (Rossi) N (Kirby) N, HL + + + (Kirby) + + + + + + (L.) + + + + + + + Stephens N + Vander Linden N + (L.) Smith N, Er + (L.) (Kirby) R + + + (L.) + + + + + + + (Kirby) + + + + + (Panzer) (Forster) Fabr. + + + + + Morawitz (continued) Apidae Andrena angustior Andrena argentata Andrena barbilabris Andrena bicolor Andrena bimaculata Andrena bucephala Andrena chrysosceles Andrena cineraria Andrena clarkella Nysson spinosus Nysson trimaculatus Oxybelus argentatus Oxybelus uniglumis Passaloecus corniger Passaloecus monilicornis Pemphredon inornata Pemphredon lethifera Pemphredon lugubris Pemphredon morio Philanthus triangulum Psenulus pallipes Rhopalum coarctatum Stigmus solskyi Tachysphex pompiliformis Trypoxylon attenuatum Trypoxylon clavicerum Trypoxylon figulus Rhopalum clavipes APPENDIX. ACULEATE HYMENOPTERA BR. J. ENT. NAT. HIST., 23: 2010 95 + Park Sutton (continued) ? + Highgate Common + + + Common Hartlebury ++ + + Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley Allioni) + + + + + + in (L.) rez + + + + + ´ (Fabr.) + + + + + + + Perkins + + + + (Kirby) + + + (Kirby) + + + + + + + (Panzer) Thomson (RDB1), HL ller) (Panzer) + + + + Pe (Seidl) + + + + + ¨ Zetterstedt R, HL + + ller Nylander + + (Kirby) N + + (L.) + + + + + + ¨ (Kirby) + (Kirby) R, Er + + + + + + (Kirby) + + + + + (Kirby) R, HL + + + + (Kirby) R, HL + + + (Kirby) R Imhoff N, HL(Kirby) + + + + + + + Nylander R, HL Fabr. N Perkins + + + + + + + (Kirby) N Smith + + + + + (Mu (Mu (continued) Andrena ovatula Andrena semilaevis Andrena scotica Andrena subopaca Andrena synadelpha Andrena tarsata Andrena nitida Andrena tibialis Anthidium manicatum Andrena wilkella Anthophora furcata Andrena labiata Andrena haemorrhoa Andrena fuscipes Andrena humilis Andrena labialis Andrena nigroaenea Andrena nigrospina Andrena fucata Andrena nigriceps Andrena coitana Andrena denticulata Andrena dorsata Andrena fulva Andrena lapponica Andrena minutula Bombus bohemicus Bombus campestris Bombus hortorum APPENDIX. ACULEATE HYMENOPTERA 96 BR. J. ENT. NAT. HIST., 23: 2010 Park Sutton (continued) + Highgate Common Common Hartlebury + ++ + Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley (Kirby) R + (Schenck) RDB3, HL + + ller) ¨ (Fabr.) + + + (L.) + + + + + + + (Thomson) R + + + (Christ) + + + + + + + Nylander + + + + (Scopoli) + + + + + + + Smith + + + ? ? Smith + + + (Mu Nylander + + + (L.) R, Er + + + + + + (L.) R + + + + + (L.) + + + + + + + (Fabr.) N, Coastal + + (L.) + + + + + + + Lepeletier + + + + + (L.) + + + + + + + (Fabr.) (Panzer) N* (R) + + (L.) + + + + + + + (Panzer) R, Er + + + + + Vogt R, HL + (Kirby) (Geoffroy) + + + + + + + (Geoffroy) R, HL + + Schenck R + + Illiger N (continued) Bombus rupestris Bombus sylvestris Bombus terrestris Bombus vestalis Chelostoma campanularum Colletes daviesanus Colletes fodiens Colletes similis Epeolus cruciger Colletes succinctus Dasypoda hirtipes Epeolus variegatus Halictus rubicundus Halictus tumulorum Hoplitis claviventris Hylaeus brevicornis Hylaeus communis Hylaeus hyalinatus Hylaeus signatus Lasioglossum brevicorne Lasioglossum albipes Bombus lapidarius Bombus lucorum Bombus magnus Bombus pascuorum Bombus pratorum Bombus ruderarius Bombus jonellus Bomus humilis APPENDIX. ACULEATE HYMENOPTERA BR. J. ENT. NAT. HIST., 23: 2010 97 Park Sutton (continued) + Highgate Common ++ Common Hartlebury Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley (Kirby) + + (Schenck) R, HL + + + + rez) + + + (Kirby) N, HL + + + ´ (Kirby) + + + rez) R, HL + + ? + (Schrank) + + + + + ´ (Pe (Scopoli) + + + + + (Kirby)(Fabr.) R, HL + + + + + + + (Kirby) + + + + (Kirby) + + + + (Schenck) + + (Schenck) R, HL + + (Pe N, HL + + + + + (Kirby) + + + + + (Zetterstedt) + + + + + ´ (Kirby) + + + (Kirby) + + + Smith + + + + (Kirby) RDB3* (no status) + + + + + + (Kirby) + + + + + + + (Kirby) R, Coastal + + (Fabr.) + + (Kirby) + + + Fabr. RDB3, HL (L.) + + (Kirby) N + + (Panzer) R ? + Brulle Panzer + + + + + + + (continued) Nomada integra Lasioglossum calceatum Lasioglosum cupromicans Nomada lathburiana Nomada marshamella Lasioglossum quadrinotatum Lasioglossum morio Lasioglossum parvulum Lasioglossum punctatissimum Lasioglossum rufitarse Lasioglossum smeathmanellum Lasioglossum villosulum Megachile ligniseca Megachile maritima Megachile versicolor Megachile willughbiella Melitta haemorrhoidalis Nomada flava Nomada flavoguttata Lasioglossum fratellum Lasioglossum lativentre Lasioglossum leucopus Lasioglossum leucozonium Lasioglossum minutissimum Melitta leporina Nomada flavopicta Nomada fulvicornis APPENDIX. ACULEATE HYMENOPTERA 98 BR. J. ENT. NAT. HIST., 23: 2010 + Park Sutton + + Highgate Common + Common Hartlebury Common Gentleshaw Devil’s Spittleful Common Barlaston/ Rough Close Common Baddesley (Kirby) + + + + + + + Hagens N + + Hagens N Thomson N, HL + + + (kirby) + + + + + + Thomson + + + + Hagens R Smith + + + + + + (L.) + + + + + + (L.) (L.) + + + + Thomson N* (no status) + + + (L.) + + + + Lepeletier + + Jurine RDB2, HL + + Hagens RDB3 Fabr. R, HL + + + + + + Fabr. + + + + (continued) (L.) + + + + Nomada rufipes Nomada signata Osmia caerulescens Nomada ruficornis Sphecodes pellucidus Sphecodes puncticeps Sphecodes reticulatus Sphecodes rubicundus Sphecodes ephippius Sphecodes ferruginatus Sphecodes geoffrellus Sphecodes gibbus Sphecodes hyalinatus Sphecodes crassus Osmia rufa Sphecodes monilicornis Sphecodes niger APPENDIX. ACULEATE HYMENOPTERA