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GRASSHOPPERS OF THE SANDLING HEATHLANDS TIM GARDINER Introduction There has been a National Vegetation Classification (NVC) system for plants in Britain for over two decades. The NVC is split into several volumes, each one dealing with specific habitats (e.g. grasslands, heathlands and mires, and woodlands and scrub). It is possible to compare survey data collecting using standardised NVC methodology with the specific plant community in the respective volume. It is then possible to identify if the particular plant community is of particular conservation interest due to its rarity. NVC plant communities may indicate specific geology and soil types, knowing this helps to target conservation management to preserve the most important plant communities. Having a system to classify grasshopper assemblages of importance may be a useful tool to identify the most valuable sites for nature conservation in Britain. In theory, it should be relatively easy to record the grasshoppers present across a range of sites in the same general habitat and then see if there are clear similarities between different types of lowland heathlands for example. Different grasshopper species have contrasting microclimatic preferences which may drive the diversity of assemblages. Short grassland and heathland swards may be unfavourable for grasshoppers due to high microclimatic temperatures (>44oC), which can lead to shade-seeking behaviour and vigorous escape responses in several grasshopper species (Uvarov 1966; Willott 1997). The optimum air temperature for development of grasshoppers in the UK is thought to be 35–40oC (Willott, 1997). Responses to microclimatic temperatures may differ between species (Gardiner & Hassall, 2009), for example, the mottled grasshopper Myrmeleotettix maculatus is a short sward specialist and its small size may be an adaptation for the high temperatures it experiences (Willott, 1997). Contrastingly, the common green grasshopper Omocestus viridulus, a long grass species in the UK, is a large insect which may overheat in short, hot grasslands/heathlands and it therefore avoids those habitats (Willott, 1997). The Sandlings are an area of coastal sands which stretch from Ipswich in Suffolk all the way north to Belton and Waveney Forest in east Norfolk. The Sandlings contain remnants of lowland heathland, approximately 1681 ha or 8% of once extensive heaths. From 1932–1983 losses of Sandling heaths were high, with 83% disappearing due mainly to afforestation (30%) and agriculture (30%). There are 42 Sandling heaths remaining, therefore, protection of these sites and their grasshopper assemblages is of paramount importance. Succession of open heathland to scrub and woodland is the current threat to the grasshoppers of open habitats, but recreational pressure, overgrazing and summer fires are also a problem. Heathlands are an outstanding habitat for Orthoptera and identifying assemblages will aid the conservation of this important resource on the Sandlings. The small-scale study described here concentrates on surveys of the Sandling heathlands in Suffolk and east Norfolk in an attempt to classify their grasshopper (Orthoptera: Acrididae) assemblages.

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Grasshopper Classification System While undertaking surveys of the grasshopper assemblages of the Sandling heathlands in Suffolk and east Norfolk in 2011, I decided to use a standardised recording method across a range of sites to collect data on the most frequent grasshopper (only Orthoptera: Acrididae noted) species in each area. Three distinct heathland areas were chosen for surveys (Dunwich, Ipswich and Waveney Forest). In all three areas, eight sites each were visited in the period July–September in 2011 (Table 1) and approximately one hour was spent at each site covering an area of c. 100 × 100 m (1 ha) recording the grasshopper species present using a combination of visual sightings and detection of stridulating males. It should be noted that the surveys at each site were samples and that species could have been overlooked; therefore, additional species may be recorded at all sites. For all of the heaths in the three areas (Dunwich, Ipswich and Waveney Forest), recording was focused in areas of bell heather Erica cinerea, heather Calluna vulgaris and stands of purple moor-grass Molinia caerulea where it occurred. Ground covered in lichens, mosses and sheep’s sorrel Rumex acetosella was also searched where it occurred, No attempt was made to record the abundance of different grasshopper species due to the difficulty in finding a suitable recording method which would accurately estimate densities in tall and short swards (Gardiner et al., 2005). Therefore, estimates of abundance would be fairly meaningless between shorter heathland swards (with C. vulgaris and mosses) and taller ones (dominated by M. caerulea). To analyse the data collected from each of the three areas, species were given a frequency value similar to that used in the NVC system for plants. The frequency value represented the number of sites that each species was recorded at as a percentage of the total number surveyed in each area (e.g. species found at 4 sites out of 8=50% frequency or III on the following scale). The frequency values were as follows: I=0–20% frequency, II=21–40%, III=41– 60%, IV=61–80% and V=81–100%. The classification of the three heathland areas was based on the two most frequent species (referred to as the dominants), with special mention made of the rarest species most likely to be of particular conservation interest. Finally, a mean species richness value was calculated for the eight surveys (equating to species richness per ha) in each area to determine the diversity of the assemblage. Results and discussion The survey revealed that the Ipswich heaths in south Suffolk were particularly species-rich (Table 2). The grasshopper assemblage of the Ipswich heaths had six species recorded, with field grasshopper Chorthippus brunneus and M. maculatus the dominant species. These two grasshoppers are insects of dry heathland on acidic soils (with abundant mosses/lichens and R. acetosella) with plenty of exposed ground (bare earth) and short swards. They can be found in high abundance in such habitats. Interestingly, O. viridulus was a rare species in the dry climate of the Ipswich heaths, probably because this insect prefers damper heathland in higher precipitation areas with tall grassland (dominated by M. caerulea) in south-east England (e.g. Epping Forest; Gardiner, 2010). Therefore, its scarcity on the Dunwich and Ipswich

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Table 1. Heathland sites visited to record grasshoppers (Orthoptera: Acrididae) in the three heathland areas of the Sandlings Area/recording site/designation Dunwich heaths Dunwich Forest – open heath no ponies Dunwich Forest – open heath ponies Dunwich Forest – pony grazed woodland Dunwich Heath – National Trust Toby’s Walks Walberswick Common Westleton Common/Heath Walberswick – New Delights Ipswich heaths Bixley Heath Foxhall Heath Martlesham Common Martlesham Heath Piper’s Vale Purdis Heath Ransomes Europark Rushmere Heath Waveney Ridge heaths Belton Common north Belton Common south Herringfleet Hills Waveney Forest disused railway line Waveney Forest felled area Waveney Forest heathland under pylons Waveney Forest wet heathland/bog Waveney Forest WWII bunkers

Grid reference No. grasshopper species TM454722 TM455723 TM467711 TM473679 TM446743 TM482749 TM454696 TM453728

6 2 1 3 2 3 7 4

TM199430 TM215444 TM245460 TM239451 TM176416 TM211425 TM206414 TM199451

5 2 2 6 3 5 4 5

TG476021 TG477019 TM468981 TG458005 TG462004 TG457005 TG457005 TG462003

4 5 5 4 3 6 2 2

heaths of Suffolk where rainfall is low is no surprise. Where this grasshopper occurs in the C. brunneus-M. maculatus (referred to as H1 assemblage in rest of paper) assemblage it will be of particular conservation interest and its localised damp and ‘cool’ habitats should be preserved. Habitats on the Ipswich heaths of particular interest for grasshoppers were the areas of short vegetation with a ‘warm’ microclimate, dominated by lichens and mosses, ideal for M. maculatus which was abundant in these situations. Of the eight sites visited, Martlesham Heath had the highest

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number of species recorded (six), while only two species were observed at Foxhall Heath and just one at Martlesham Common. The acid grassland at Foxhall Heath was situated on the approach to the Stock Car Stadium and must be under severe pressure from recreational usage (leading to trampling of swards) before races. Despite this, the uniformly short swards still have value for an extremely localised population of M. maculatus, which is associated with a patch of mossy ground where road planings have been laid as a surfacing material. The species-poor assemblage at Martlesham Common may be due to a fire in 1995 which was thought to have eradicated the Nationally Scarce silver-studded blue Plebeius argus butterfly from the site. At Purdis Heath, five species were recorded. Management on this heath for P. argus in the form of Birch Betula spp. and Gorse Ulex europaeus clearance and creation of disturbed, open ground is likely to be beneficial for Orthoptera which could suffer from unmanaged encroachment of scrub. Certain grasshoppers favour disturbed heathy ground at an early stage of succession (C. brunneus and M. maculatus) similar to P. argus (Ravenscroft, 2009). These grasshopper species are ‘disturbance-dependent’ (Gardiner & Benton, 2009). In the areas managed for the butterfly at Purdis Heath (one adult was even seen in flight during the grasshopper survey at this site), M. maculatus was found utilising the bare earth and sparse vegetation. There is an interesting contrast between the grasshopper assemblage diversity at the three Ipswich sites (Martlesham Heath, Purdis Heath and Ransomes Europark) where P. argus has been recorded in recent years (5 species/site) compared to those where the butterfly has not (3∙4 species/site). This suggests that where P. argus occurs there are particularly species-rich grasshopper assemblages. The stripe-winged grasshopper Stenobothrus lineatus was associated with acid grassland and ant hills on the Ipswich heaths, a habitat it seems to require in other counties (e.g. Essex and Norfolk). Fortunately, three of the Ipswich heaths surveyed are now protected by designation as Sites of Special Scientific Interest (SSSI) and are actively managed to maintain and increase their biodiversity. This is particularly important as the heathlands of Ipswich are under immense pressure from development and much habitat has been lost due to industrial and residential developments since the 1940s. The legal protection should ensure no further losses of biologically important heathland in Ipswich, although the scraps of dry heathland still remaining on the Ransomes Europark development (with E. cinerea, R. acetosella and plentiful mosses), closely guarded and fenced off from the public, may one day disappear under concrete and tarmac. Ironically, around the edges of one of these fenced off remnants, scattered road planings formed suitable sparsely vegetated habitat for C. brunneus and M. maculatus. One such heathland remnant, partially designated as a County Wildlife Site (CWS), has had P. argus recorded (Ravenscroft, 2009; Parker, 2012) and this has being taken into consideration in planning applications for the site. It must be hoped that further development can be resisted on these biologically important remnants of the Sandlings.

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The Dunwich heaths, although dominated by the same two grasshopper species (C. brunneus-M. maculatus), were less species-rich overall than the Ipswich heaths (Table 2). The Nationally Scarce woodland grasshopper Omocestus rufipes was present at two ungrazed sites near Dunwich Forest (Gardiner, 2012a) and is very rarely found to the north of the Sandling heaths in the UK. Therefore, where this species occurs in Dunwich Forest it should be the subject of targeted conservation management. It was absent from heathland heavily grazed by Dartmoor ponies due to the lack of structural diversity in the sward which was uniformly short with few patches of taller grassland or Calluna (Gardiner, 2012a). The assemblages in pony grazed woodland (one species) and heathland (two species) at Dunwich Forest were very species-poor, compared to ungrazed vegetation (six species including M. maculatus, O. rufipes and S. lineatus). Recreational pressure on heathland was a serious issue at Toby’s Walks where uniformly short acid grassland and Calluna swards with plenty of bare earth have become established by the wear and tear of walkers’ feet. However, C. brunneus and M. maculatus, two insects of early successional stages with pioneer vegetation and bare earth (Gardiner & Benton, 2009), appeared to be common at Toby’s Walks suggesting that trampling of heathland can be beneficial to these two grasshoppers, but detrimental to other species of taller, damper swards (e.g. meadow grasshopper Chorthippus parallelus and O. viridulus). Somewhat surprisingly, the dry heathland of Dunwich Heath owned and managed by the National Trust was fairly species-poor (only three species), with S. lineatus, O. rufipes and O. viridulus all absent. The most numerous grasshoppers at Dunwich Heath were C. brunneus and M. maculatus, perhaps indicating the lack of wet heathland which species such as O. viridulus require. The outstanding site in the entire survey was Westleton Heath (all seven species recorded), a superb example of Sandling heathland with patches of dry Calluna (with C. brunneus and M. maculatus) and Molinia tussocks (with O. rufipes and O. viridulus) interspersed with sporadic Betula trees. This habitat is as close to the New Forest heathlands as you can get in eastern England, and not surprisingly it is home to one of the last remaining O. rufipes populations in East Anglia. Indeed, inspection of Marshall & Haes (1988) shows that the total of seven grasshopper species at Westleton Heath (c. 54% of British total of 13 grasshopper species) is notable in a national context, comparing favourably with the best areas for acridids in the UK such as the Dorset heathlands (nine species), New Forest (eight species) and the Purbeck Hills (seven species). The diverse mixture of vegetation stands and structure (e.g. short and tall vegetation in close proximity) managed by light cattle grazing make it an extremely important site for grasshoppers. Once again, there was a relationship between the occurrence of P. argus on sites on the Dunwich Heaths (near New Delights (Parker, 2012) and Westleton Common/Heath) and high species richness of grasshoppers (with P. argus: 5∙5 species/site, without: 2∙8). Therefore, it really does seem that management to benefit this rare butterfly is favourable for grasshoppers on the Dunwich and Ipswich Sandlings and can promote high diversity of acridids.

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In contrast to the more threatened Ipswich heaths, most of the Dunwich heaths surveyed (six sites) are protected by a SSSI designation, but also by a Special Area of Conservation (SAC), Special Protection Area (SPA) and Ramsar Site, which should ensure the continued maintenance of the open acid grassland and dry and wet heathland required by Orthoptera. The assemblages of the relatively dry heathlands of the Dunwich and Ipswich heaths contrasted with the rarer, more distinctive assemblage of the Waveney Forest heaths situated in east Norfolk. The main survey sites were located in and around Waveney Forest. Chorthippus brunneus and O. viridulus were the two dominant species. This is a rather unusual combination of species, as the former indicates dry heathland, while the latter wet heathland (Marshall & Haes, 1988). Therefore, the presence of both species at the same sites indicates an intricate mixture of dry and wet heathland reflecting the complex geology of the gravel ridge of the Waveney Forest area (Gardiner, 2012b). The main survey areas were stands of M. caerulea dominant over peaty soils, although forestry operations in Waveney Forest (e.g. felling of trees and transport of cut timber) had led to the establishment of patches of bare earth among and beside the Molinia tussocks, which were suitable for small populations of C. brunneus. It is believed that the latter species can tolerate a wide range of sward heights (Willott, 1997) and can be found in the taller grassland of wet heathland as a consequence. Dry heathland was also very close to the stands of Molinia, meaning that C. brunneus individuals may have strayed onto the wet heathland where exposed soil was present. Wet heathland vegetation predominates at many sites in the Forest which is why M. maculatus and S. lineatus were not as frequent as in the dry heathland assemblages of the Dunwich and Ipswich heaths (Table 2). The quite distinct assemblage of interconnected dry and wet heathland is referred to in this study as C. brunneus-O. viridulus (mentioned as H2 in rest of paper). In south-east England this assemblage is probably restricted to certain areas such as east Norfolk where C. parallelus is not as widespread as in the rest of the UK (Richmond, 2001). Wet heathland usually has C. parallelus as a key component, but in some areas where it is not widespread it may be replaced by other grasshoppers such as C. brunneus in Waveney Forest (Gardiner, 2013). From a microclimatic perspective, ‘cool’ wet heathland swards dominated by Molinia are favourable for O. viridulus, but not M. maculatus which is a dry heathland species, associated with ‘warm’ swards of early successional stages with plentiful bare earth, lichens and mosses. Management of the heathlands may also determine their grasshopper diversity. For example, heavy trampling by walkers and pony grazing on the Dunwich heaths effectively produces a sward composed entirely of the early successional grasshoppers C. brunneus and M. maculatus, but eradicates all the species of taller vegetation (e.g. O. viridulus) and those which require structural diversity such as the scarce O. rufipes. Urgent conservation action is needed to protect the localised areas of Dunwich Forest and Westleton Heath which support O. rufipes. Overgrazing by ponies produces a short heathland sward

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and must be tackled at these sites to promote the development of structurally diverse vegetation with short and tall patches which can support up to seven grasshopper species. It appears that management to increase P. argus populations may have the added benefit of conserving diverse grasshopper assemblages. The Dunwich and Ipswich heaths are often designated as SSSIs, so the long-term survival of the grasshopper assemblages should be ensured. However, the dry and wet heaths of the Waveney Forest area have no such protection, although small areas (e.g. Belton Common) are CWS. Overall, on the Sandlings heaths, sites protected by SSSI designations had higher grasshopper species richness (4∙6 species/site) than those designated as CWS or without any form of protection (3∙3 species/site). Therefore, the most valuable heaths for grasshoppers will probably be conserved for future generations to enjoy and should receive favourable management (often to benefit P. argus). Recently proposals for large-scale gravel extraction have been put forward for the Forest; fortunately these now appear to have been successfully defeated by a well organised local protest (Gardiner, 2012b). Because the Forest is privately owned, management for nature conservation is not undertaken and scrub encroachment by Betula spp. is now a serious problem for grasshoppers of the open areas. Only 6% of the Forest is open heathland, planting of conifers since the 1940s has reduced its extent massively. Fortunately, swathes of conifers are now being felled as they have reached maturity; the open areas this has created are favourable for early successional grasshoppers such as M. maculatus, until such time when the replanted conifers shade out the woodland floor once again. Hopefully, grasshoppers can follow the clear-felling around the Forest, taking advantage of the warm microclimate established as the trees are felled. Other grasshoppers which should benefit from clear-felling are C. brunneus and O. viridulus (the latter where Molinia tussocks regenerate). A new method for classifying grasshopper assemblages It is necessary to trial the classification system in other areas of heathland throughout the south-east of England to see if it is possible to determine additional assemblages of conservation importance. To check the classification system against another area with historic records using roughly the same method, I compared the 2011 Sandlings data against that collected during surveys of eight sites in July 2010 on wet heathlands in Epping Forest dominated by M. caerulea with occasional cross-leaved heath Erica tetralix. These wet, grassy heathlands had the lowest species richness of all the East Anglian sites, with only five grasshopper species recorded (mean species richness = 3∙1 species/site, range 2/4). The dominant species were C. parallelus and O. viridulus (both had a frequency of V), both insects of taller grassland and heathland on wetter soils. There was a low frequency (I) of S. lineatus, typically a species of drier habitats but increasing its range in the Forest after the first record for the county in 2009 on Whitehall Plain (Wilde, 2009). There was a complete absence of M. maculatus, which is probably

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extinct in Epping Forest due to the decline in grazing management in the 1900s and consequent scarcity of early successional habitats in the tall and tussocky Molina dominated swards (Gardiner, 2010). This assemblage (referred to as H3) is therefore quite a distinct one from the dry heath (H1) of the Dunwich and Ipswich areas and the dry/wet mosaic heath (H2) of Waveney Forest previously described. The other two species frequently recorded in Epping Forest were C. albomarginatus (frequency IV) and C. brunneus (frequency III). It must be noted that the C. parallelus-O. viridulus wet heath assemblage may probably be different to that of quaking bogs found in areas such as the New Forest. Therefore, so far three distinct grasshopper assemblages have been identified indicative of a particular type of heathland: dry heath (C. brunneus-M. maculatus), dry/wet heath mosaic (C. brunneus-O. viridulus) and wet heath (C. parallelus-O. viridulus). The wet heath classification of Epping Forest was compared with other survey data from Culm Grassland in north Devon, south-west England. In August 2004, seven sites were visited by the author at Dunsdon National Nature Reserve and Quoditch Moor. At these sites, M. caerulea stands predominated with patches of E. tetralix and the dominant grasshoppers were C. parallelus and O. viridulus (both had a frequency of V), with C. brunneus the only other species recorded (frequency of III). The mean species richness (2∙3 species/site, range 1–3) was extremely low, as it was in stands of M. caerulea in Epping Forest. Therefore, it would appear that the species-poor wet heathland assemblage of the north Devon Culm Grasslands conformed to the wet heath assemblage of Epping Forest. This indicates that it may be possible to classify specific heathland assemblages in other regions of the UK by their dominant grasshopper species. It should be noted that the wet heathland assemblage (and species such as O. viridulus) is of more conservation importance in the drier climate of eastern England where it is rarer, than it will be in the wetter west of England where the habitat it occupies is much more frequent. Grasshopper species richness is likely to be highest in dry heath (H1=3∙5– 4∙0) and the dry/wet heath (H2) mosaic (3∙9), both relatively species-rich assemblages, compared to the species-poor wet heath (H3) stands of M. caerulea (2∙3–3∙1). Therefore, some interesting assemblage traits are beginning to appear from the standardised study of heathlands on the Sandlings, but also further afield in Epping Forest and the Culm Grasslands. Presentation of results for the assemblage of a particular geographical area or heathland stand should be similar to that in Table 2 enabling comparison with the current survey results. Inclusion of species richness and frequency values is essential to determine the structure of the assemblage and the two dominant species. Where three or four species have the same overall frequency for an area, a subjective assessment should be made of the respective abundances to determine the co-dominants. Rarer species of note should be highlighted for each geographical area. Assemblages of heathland in other regions of the UK should be published to determine the whole range of grasshopper assemblages possible. A separate code should be assigned to the new assemblage (e.g. H4) to differentiate it from those identified in this paper.

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A particularly good area to undertake a further trial of the system in East Anglia would be the Breckland heaths in Thetford Forest. The valley mires of the New Forest also warrant investigation and may hold a rare type of assemblage which is likely to be of high conservation importance due to the presence of the large marsh grasshopper Stethophyma grossum, an Endangered Species listed in the British Red Data Book. Preliminary surveys from six New Forest mires in 2012 indicate a C. parallelus-S.grossum assemblage with a low overall species richness. Table 2. Frequency of each grasshopper species in the three heathland areas of the Sandlings Species Chorthippus albomarginatus Chorthippus brunneus Chorthippus parallelus Myrmeleotettix maculatus Omocestus rufipes Omocestus viridulus Stenobothrus lineatus Mean species richness/ha (min/max) Predominant habitats Grasshopper Classification System code

Dunwich

Ipswich

Waveney

II V II V II I III

IV V IV V I III

II V II IV V I

3∙5 (1/7) 4∙0 (1/6) Dry heath Dry heath H1

H1

3∙9 (2/6) Dry/wet heath H2

Frequency: I=1–20% of sites, II=21–40%, III=41–60%, IV=61–80%, V=81–100% References Gardiner T. (2010). Precipitation and habitat degradation influence the occurrence of the common green grasshopper Omocestus viridulus in southeastern England. Journal of Orthoptera Research 19: 315–326. Gardiner T. (2012a). Rediscovery of the woodland grasshopper Omocestus rufipes in East Suffolk. Transactions of the Suffolk Naturalists’ Society 47: 53–54. Gardiner T. (2012b). The Battle to Save a Norfolk Forest: The story so far…. Country-Side 33 (4): 16–20. Gardiner T. (2013). Grasshoppers (Orthoptera) of Great Yarmouth. Transactions of the Norfolk & Norwich Naturalists’ Society 45 (1): 84–94. Gardiner T. & Benton T. (2009). Grasshoppers and bush-crickets (Orthoptera) of military training grounds near Colchester. Entomologist’s Record & Journal of Variation 121: 167–171. Gardiner T. & Hassall M. (2009). Does microclimate affect grasshopper populations after cutting of hay in improved grassland? Journal of Insect Conservation 13: 97–102.

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Gardiner T., Hill J. & Chesmore D. (2005). Review of the methods frequently used to estimate the abundance of Orthoptera in grassland ecosystems. Journal of Insect Conservation 9: 151–173. Marshall J. A. & Haes E. C. M. (1988). Grasshoppers and Allied Insects of Great Britain and Ireland. Harley Books, Colchester. Parker R. (2012). 2010 Butterfly Report. Transactions of the Suffolk Naturalists’ Society 47: 80–92. Ravenscroft N. (2009). The Silver-studded Blue on the Sandlings, Report to Butterfly Conservation (Suffolk Branch). Butterfly Conservation (Suffolk), Bury St. Edmunds. Richmond D. (2001). Grasshoppers and Allied Insects of Norfolk. Norfolk & Norwich Naturalists ‘ Society, Norwich. Wilde I. (2009). The Stripe-winged Grasshopper Stenobothrus lineatus (Panzer, 1796) (Orthoptera: Gomphocerinae) new to Essex. Essex Naturalist (New Series) 26: 61–62. Willott S. J. (1997). Thermoregulation in four species of British grasshoppers (Orthoptera: Acrididae). Functional Ecology 11: 705–713. Uvarov B. P. (1966). Grasshoppers and Locusts, Vol 1. Cambridge University Press, Cambridge. Tim Gardiner Environment Agency, Iceni House, Cobham Road, Ipswich, Suffolk IP3 9JD timgardiner134@btinternet.com

Trans. Suffolk Nat. Soc. 49 (2013)

GRASSHOPPERS OF THE SANDLING HEATHLANDS  

Tim Gardiner

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