Review Article Journal of Orthoptera Research 2018, 27(1): 91-96 The effects of grazing and mowing on large marsh grasshopper, Stethophyma grossum (Orthoptera: Acrididae), populations in Western Europe: a review JAc@ul MILLeR', Tim GARDINER? 1 RSPB, Stalham House, 65 Thorpe Road, Norwich, UK. 2 Environment Agency, Iceni House, Cobham Road, Ipswich, Suffolk, UK. Corresponding author: Jacqui Miller (jacqui.miller@rspb.org.uk) Academic editor: Corinna S. Bazelet | Received 25 January 2018 | Accepted 25 April 2018 | Published 12 June 2018 http://zoobank.org/787ED432-BEBC-4A66-8651-DB6A2BD35196 Citation: Miller J, Gardiner T (2018) The effects of grazing and mowing on large marsh grasshopper, Stethophyma grossum (Orthoptera: Acrididae), populations in Western Europe: a review. Journal of Orthoptera Research 27(1): 91-96. https://doi.org/10.3897/jor.27.23835 Abstract The large marsh grasshopper, Stethophyma grossum L. (Orthoptera: Acrididae), has undergone a significant range contraction in the UK and is now restricted to the bogs and mires of the New Forest and Dorset Heaths. In other parts of Western Europe, the species makes use of a wider range of wetland habitat types. Traditionally, many of these habitats would be man- aged through low intensity grazing, mowing, or both, and these measures are now often employed in the conservation management of wet grassland habitats. This paper reviews the effects of mowing and grazing on S. gros- sum populations, through looking at the potential impacts (both positive and negative) on different life stages of the grasshopper. Both techniques are valuable in the maintenance of an open and varied vegetation structure which is known to benefit S. grossum in all its life stages. However, grazing on very wet sites or at high intensity can result in trampling of vegetation and S. grossum eggs, and mowing which is too frequent may negatively affect populations through repeated losses of nymphs. Recommendations are given regarding the suitability of mowing and grazing for different habitats and intensity of management to generate the required vegetation structure. Measures are also outlined, such as the provision of unmown or ungrazed refuge areas, which can help reduce negative effects. Key words adults, biodiversity conservation, bog, eggs, grassland, management, mire, nymphs, vegetation structure, wetland Introduction Grazing and mowing exert important influences on vegetation structure and are therefore key factors affecting grasshopper popu- lations (Clarke 1948, Gardiner et al. 2002, Humbert et al. 2009, Kenyeres and Szentirmai 2017). Rare and localized species, such as the large marsh grasshopper, Stethophyma grossum L. (Orthoptera: Acrididae), have very specific micro-habitat requirements which can be influenced by grazing and mowing. In the UK S. grossum is a priority species under the NERC Act 2006 and has a GB IUCN status of Near Threatened (Sutton 2015). It has undergone the largest range contraction of all the UK Orthoptera between the 1980s and 2000s (Beckmann et al. 2015), and is currently con- fined to the Dorset Heaths and New Forest. In Europe, it is locally distributed with an IUCN status of Least Concern (Hochkirch et al. 2016), however, in Switzerland and Austria it is listed as Vulner- able (Berg et al. 2005, Monnerat et al. 2007) and in Denmark it is considered Near Threatened (Wind and Pihl 2010). It is the aim of this paper to describe what is known about the links between the life cycle and habitat requirements of S. grossum and provide a dis- cussion of the benefits and disadvantages of mowing and grazing for the management of this species in Western Europe. Stethophyma grossum distribution and life history Stethophyma grossum (Figs 1, 2) is locally distributed across Eu- rope and found from Ireland in the west, northern Spain and Italy in the south, east to Siberia and north as far as parts of Scandina- via (JNCC 2010, Benton 2012). In the UK, its former distribution was in suitable habitat south of a line from the Bristol Channel to the Wash, although it has experienced a sharp contraction in its range and is now confined to the Sphagnum-dominated bogs and mires of east Dorset and the New Forest (Benton 2012). Popula- tions are thought to have been relatively stable in the New Forest over the last 20 years, and there is potential for the species to ben- efit from mire restoration projects underway in the Forest (Harvey and Brock 2017). In Ireland, the species is found primarily in bogs and mires, with some records from more grassy habitats, and it is locally distributed across the west, south-west and central parts of the country (Sutton 2017). S. grossum is herbivorous, feeding on the stems and seed heads of grasses, rushes and sedges (Benton 2012). Adults can be seen from late July through to October or even early November (Haes and Harding 1997, Benton 2012). They lay up to 14 eggs in the late summer in an elongated pod at the base of grass stems (Ben- ton 2012). The nymphs usually emerge in late May and early June the following year and pass through four or five instars before reaching the adult stage in late summer (Evans and Edmondson 2007, Benton 2012). JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1) ‘oO NO "eee es «oo o> 4 w! Fig. 2. Male Stethophyma grossum; credit P. Brock. Habitat types used by S. grossum In the UK, S. grossum is typically found on quaking acid bogs with purple moor-grass, Molinia caerulea (L.) Moench, bog myrtle, Myrica gale L., cross-leaved heath, Erica tetralix L., broad-leaved cot- ton grass, Eriophorum latifolium Hoppe, and white beak-sedge, Rhyn- chospora alba (L.) Vahl (Haes and Harding 1997, Edwards 2002, J. MILLER AND T. GARDINER Fig. 3. Sphagnum-dominated mire in the New Forest, UK; habitat for Stethophyma grossum; credit T. Gardiner. Fig. 4. Metrioptera brachyptera; credit T. Gardiner. Benton 2012). In the New Forest in southern England (Fig. 3), it shows a preference for Sphagnum-dominated mires with open wa- ter and wet areas indicated by cotton grass, and often coexists with the bog bush-cricket, Metrioptera brachyptera L. (Orthoptera: Tetti- goniidae; Fig. 4) (Benton 2012, Harvey and Brock 2017). Similarly, Cheesman and Brown (1998) report that S. grossum occurrence shows a positive correlation with area of surface water, cover of Sphagnum and white beak-sedge and a negative correlation with ericoids and sub-shrubs. The species typically inhabits the wettest parts of such habitats (Ragge 1965), and has even been observed swimming across bog pools in the New Forest (Gardiner 2013). Its former distribution in the UK and current distribution in the rest of Western Europe shows a wider habitat usage, including areas of fenland, moorland, wet meadow and riverside (Benton 2012). Lucas (1920) noted a record of the species from Norfolk in 1892 occurring in tall rank grass close to a river bank and Marshall and Haes (1988) suggested that the few remaining fenland popu- lations in England at that time were found in very wet conditions among sedge and grass tussocks. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1) J. MILLER AND T. GARDINER 93 Malkus (1997) noted that vegetation structure appeared to be particularly important in determining distribution of S. grossum, with nymphs being predominantly found in areas with patchy and medium-high vegetation. An open habitat structure (determined by vegetation height and density) is thought to be beneficial in allow- ing sufficient warming of the ground and the base of the vegetation to promote egg development and hatching (Malkus 1997, Marzel- li 1997, Maas et al. 2002). A study by Krause (1996) in Germany found that tufted hair grass, Deschampsia cespitosa (L.) P. Beauv., held high densities of early instar nymphs and postulated that the growth form of this plant was favorable at the time of hatching, being lower and less dense than other vegetation in the study area. Decleer et al. (2000) and Thorens and Nadig (1997) also recognize a link be- tween periodic/winter flooding and S. grossum occurrence, which may be due to the high humidity requirements of S. grossum eggs and their sensitivity to dehydration (Detzel 1998, Maas et al. 2002). Table 1 summarizes the general habitat types currently used by S. grossum in Western Europe. A wide variety of wet habitats are used, some of which will provide the required vegetation structure through management by mowing and/or grazing. Effects of mowing and grazing on life stages of S. grossum The traditional management of wet hay meadows and flood- plain grasslands in Western and Central Europe centered on hay cutting and the grazing of livestock. Many wet areas were grazed by livestock at low intensities. This was sometimes combined with cutting for hay, with one early cut followed by grazing of the rem- nant sward. Alternatively, on some sites, hay cutting was carried out once or twice a year, typically in May-June and/or August-Septem- ber (Grootjans and Verbeek 2002, Kenyeres and Szentirmai 2017). The wet heath, mire and bog habitats of S. grossum in the UK and Ireland typically have a naturally open and patchy vegetation struc- ture with areas of open water. The wettest parts of these habitats are not suitable for management by mowing or grazing, either in terms of the potential impacts on the habitat, or safety and accessibility for animals and machinery. Around the drier margins of these hab- Table 1. Habitats of S. grossum in Western Europe. itats, low intensity grazing by ponies or cattle during the summer may be used to help reduce the dominance of purple moor grass and reduce encroachment of scrub (Symes and Day 2003, Lake and Underhill-Day 2004, Groome and Shaw 2015). However, grazing of the wettest areas (mires or bogs) can be detrimental through trampling damage, particularly to bog mosses, and the creation of a more homogeneous vegetation structure (Symes and Day 2003, Groome and Shaw 2015). In the New Forest, Pinchen and Ward (2010) attribute a general decline in Orthoptera to increased graz- ing pressure since the 1960s, with trampling and changes to vegeta- tion structure likely to negatively affect many invertebrate species. While mires and bogs are less likely to be affected by overgrazing due to inaccessibility of the habitat, the effects of heavy grazing pressure were observed at two S. grossum sites in the New Forest during a recent survey (Harvey and Brock 2017). In wet grassland habitats, sensitive management by mowing and/or grazing is considered beneficial overall to S. grossum. The following section discusses considerations relating to mowing and grazing of wet grassland habitats and the requirements and char- acteristics of S. grossum eggs, nymphs and adults. Eggs.—S. grossum eggs require high humidity levels for successful development and are very sensitive to dehydration (Detzel 1998, Maas et al. 2002). Because of this, soils which are saturated or flooded during the winter are preferred (Malkus 1997). While S. grossum has relatively low temperature requirements compared to other Orthoptera (Marzelli 1997), a sufficiently open habitat structure will promote egg development and hatching (Malkus 1997, Marzelli 1997, Maas et al. 2002). Grazing while S. grossum is at the egg stage may result in the direct destruction of eggs by trampling, particularly on the wettest sites (Malkus 1997), but grazing or mowing of less wet sites can help provide the necessary open vegetation structure if carried out at low intensity and avoiding very wet areas. Nymphs.—The distribution of early instar nymphs is thought to be a product of the female choice of habitat for oviposition, as References Kleukers et al. (2004), Bakker et al. (2015) Decleer et al. (2000), Sardet et al. (2015) Country Habitat types Wet grasslands and meadows, floodplains, ditches and margins of waterbodies, fens, Netherlands swamp, wet heath. . Wet grasslands and meadows, swamp, bogs, ditches, wet heath. Land that is wet in Belgium : winter. Bette Wetlands: marshes, reedbeds, flooded meadows, peat bogs, ditches. In the Alps, up to 2400-2700 m in altitude. Luxembourg Wet meadows, marshes, peat bogs, ditches. saan Near open water or periodically flooded vegetation, wet meadows and pasture, peat bogs, ditches. Up to 2450-2700 m in altitude. Austria Peat bogs, fens, floodplains, ridges of raised bogs. Germany Marshes, edges of lakes, streams and ditches, wet meadows. Up to 1300 m in altitude. Northern Italy Lake margins, swamps, alpine fens, wet meadows. Northern Spain Wet peaty meadows, peat bog, wet mown meadows, margins of ponds and rivers. England, UK Bogs and mires in the Dorset and New Forest heaths. eee Mire, wet heath, blanket and raised bogs, Molinia-dominated grassland. By rivers and lakes. Denmark Raised bogs, wet meadows, nutrient-poor fen. Fennoscandia Bogs, meadows, by lakes and streams. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1) Voisin (2003), Sardet et al. (2015) Sardet et al. (2015) Thorens and Nadig (1997), Sardet et al. (2015) Ortner and Lechner (2015) Detzel (1998), Maas et al. (2002), Fischer et al. (2016) Galvagni (2001), Fontana and Kleukers (2002), Kranebitter (2008) Liiders (2009) Haes and Harding (1997), Benton (2012) Benton (2012), Sutton et al. (2017) Hansen and Jorgensen (2010) Holst (1986) 94 young nymphs have limited mobility and therefore do not tend to disperse from their hatching location (Marzelli 1997). Malkus (1997) found that patchy vegetation with a heterogeneous struc- ture was preferred by nymphs. As above, low intensity grazing may provide the necessary diversity in sward structure. Mowing can have a significant effect on the density of nymphs. If mowing takes place during the early summer, the density of early instar nymphs is likely to drop significantly afterwards (Krause 1996, Malkus 1997, Marzelli 1997, Detzel 1998). Due to the limited mo- bility of young nymphs, they are not able to take evasive action, and may either be directly killed, removed with the hay crop or made more vulnerable to dehydration and predation (Krause 1996, Malkus 1997). Malkus (1997) observed a collapse in nymph numbers after mowing in mid-June, however, after 1-2 weeks, numbers recovered as further hatching occurred, possibly promoted by the increased levels of solar radiation reaching the ground. Krause (1996) noted detri- mental effects on populations affected by mowing at an early stage in nymph development. Later instar nymphs may be more able to escape mowed areas — Krause (1996) noted an increase in late instar nymph densities around ditch edges following mowing. Adults.—Adult S. grossum also tend to be found in locations with relatively high soil moisture levels, perhaps due to their need to ovi- posit in wetter areas. Sonneck et al. (2008) propose that adults also benefit from a heterogeneous vegetation structure as this allows the adults to withstand fluctuating temperatures. Similarly, a variety of soil moisture levels within a site may allow adults to cope with vary- ing weather conditions (Detzel 1998, Kleukers et al. 2004). Malkus (1997) reports that mowing later in the summer (mid- July onwards) tends to displace adult S. grossum to neighboring areas until the vegetation regrows, when repopulation will occur. Grazing during this period has similar effects. Population effects as a result of mowing are unlikely unless the mowing is too frequent, takes place in cool weather when the grasshoppers are less active and therefore less able to take evading action, or where unmown refuge areas are not available (Malkus 1997). Malkus (1997) also observed adults flying up in front of a mower and moving to the as yet unmown center of the field. It is therefore possible that S. gros- sum could benefit from mowing that works from the inside of the field outwards (as is sometimes employed for certain bird species, e.g. corncrake Crex crex L.), as animals may then be more likely to reach safe habitat outside the mown area. Recommendations relating to mowing and grazing for S. grossum conservation The following recommendations are derived from the studies of S. grossum populations in Western Europe and are relevant to the management of wet grasslands including wet meadows and pasture, floodplain grassland and fens. As discussed above, S. grossum is currently only found in valley mires and bogs in the UK (and predominantly so in Ireland), therefore many of these recommendations will not be directly applicable to UK and most Irish populations. In mire and bog habitats, management should focus on protecting sites from activities likely to cause drying, although removal of encroaching scrub and/or management of dominant grasses or bog myrtle may occasionally be required on the drier margins of such sites. Mowing.—A sensitive mowing regime can be beneficial for the management of wet meadows for S. grossum (Krause 1996, S6- rens 1996, Marzelli 1997, Malkus 1997, Detzel 1998). While it J. MILLER AND T. GARDINER may cause short-term reductions in the numbers of grasshoppers (particularly early stage nymphs), if carried out with regard to their lifecycle, careful mowing can have positive effects on egg and nymph development by maintaining a more open vegeta- tion structure, thus raising ground temperatures. Insufficient, ir- regular, or mowing only in the late summer may have negative effects through matting of the turf (Malkus 1997, Detzel 1998). However, there are some differences of opinion as to the opti- mum time for mowing. Krause (1996) recommends one late cut in August, by which point most individuals should be adults and able to move to an adjacent area. Marzelli (1997) recommends two cuts — one at the beginning of June before the eggs hatch and one in mid-September after oviposition. She also notes that mowing in July was particularly damaging to populations. Mal- kus (1997) recommends that mowing should take place once and, at most, twice a year. He points out that the timing may also need to take other grassland species into account — if amphib- ians or ground-nesting birds are present, early cuts should not take place before mid/end June and the late cut should be after mid-September. It would therefore appear that wet grasslands managed for S. grossum should be cut at least once (though no more than twice) a year, depending on site-specific habitat needs, and with the aim of avoiding the vulnerable early nymph stage. If an early cut is required, ideally this should be before S. grossum has hatched, but the needs of other species present should also be considered. If a late summer cut is required, this should be after mid-September when most of the egg-laying is complete. In order to minimize mortality of grasshoppers (and other invertebrates), the use of a bar mower (rather than a rotary or flail mower) set to a minimum height of 10 cm is recommended (Humbert et al. 2009, Kenyeres and Szentirmai 2017). Malkus (1997) also makes further recom- mendations to reduce mortality of S. grossum during mowing: mowing should only take place in warm, sunny weather, to al- low grasshoppers to escape; retain the hay on the surface for a few days following the cut, again to allow grasshoppers to escape; and unmown areas should be retained close to the mown area, to provide a refuge. Grazing.—Low intensity grazing is a useful method for managing vegetation height and density and tends to create a more varied vegetation structure than mowing alone (Lake and Underhill- Day 2004). Grazing can also help prevent scrub encroachment and reduce cover of dominant species (Symes and Day 2003, Lake and Underhill-Day 2004, Groome and Shaw 2015), thus helping to maintain the open vegetation structure required by S. grossum. As well as having similar displacement effects to mow- ing, grazing has the potential to cause damage to habitats and destruction of S. grossum eggs through trampling (Malkus 1997, Groome and Shaw 2015). It is therefore important to select an appropriate livestock type and stocking rate for the habitat type; ponies will tend to create a more homogeneous sward than cat- tle (particularly if grazed at high stocking rates), although cattle may be more likely to cause trampling damage (English Nature 2005). Malkus (1997) recommended that grazing in general should be carried out at a low stocking density and on a tempo- rary basis and avoided completely on very wet habitats due to the risk of trampling damage. The potential negative effects of displacement of grasshoppers can be reduced by the retention of ungrazed refuge areas. A summary of the advantages and disadvantages of mowing and grazing are presented in Table 2. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1) J. MILLER AND T. GARDINER 95 Table 2. Advantages and disadvantages of mowing and grazing for S. grossum. Mowing Creates an open sward structure. a Restricts scrub encroachment. Mortality of nymphs. Disadvantages Displacement of adults. Potential effects on other species (e.g. ground-nesting birds). Conclusions The following recommendations for mowing and grazing as part of the management of wet grassland habitats occupied by S. grossum can be derived from this review: - Grazing and/or mowing (dependent on habitat type) are valuable management techniques for the maintenance of the open and varied vegetation structure required by S. gros- sum. - Grazing should be at a low stocking density (and, where nec- essary, for a limited time-period), and nearby ungrazed refuge areas should be maintained. — Grazing of very wet areas should be avoided. — Wet grasslands should be cut once a year, or twice at the most, depending on the vegetation type. — Ifearly mowing is used, this should be before the main hatch- ing period from mid-June onwards where possible (depend- ing on the needs of other species) and late mowing should be after the main oviposition period from mid-September onwards. Grazing Creates a varied sward structure. Reduces cover of dominant grasses. Restricts scrub encroachment. Poaching of wet habitats. Displacement of adults. Trampling of eggs. Overgrazing possible. Potential effects on other species (e.g. ground-nesting birds). — Use a bar mower set to a minimum height of 10 cm to mini- mize mortality. - Grasshoppers should be allowed to escape mowing by carrying out operations only on warm, sunny days when grasshoppers are active, retaining unmown refuge areas nearby and leaving the hay crop on the surface for a few days before removal. The following recommendations relate to the management of wet heath, mire and bog habitats, such as those used by S. grossum in the UK and Ireland: — Protect sites from activities likely to cause drying of habitats. — Grazing (and mowing) should be avoided in the wettest areas, particularly in mires and bogs. — If necessary, low intensity grazing could be used on the drier margins of such sites during the summer months to reduce dominance by grasses or encroachment of scrub. Recommended management measures for habitat types used by S. grossum in the UK and Western Europe are summarized in Table 3. Table 3. Appropriate management options for S. grossum in Western Europe. Habitat Management Alpine pasture Light grazing’ Ditch banks Mowing Fen Sedge cutting Mire/bog* Avoid grazing Reedbed Reed cutting Wet heath* Light grazing Wet grassland Mowing "Typical stocking density: 0.1 cows/ha (Homburger et al. 2015). *Only habitats left for S. grossum in the UK (Dorset, New Forest). Acknowledgements Many thanks to Paul Brock for the use of his photographs, and to the following people who provided literature sources: Bjorn Beckmann, Paul Brock, Bryan Edwards and Martin Harvey. References Bakker W, Bouwman J, Brekelmans F, Colijn E, Felix R, Grutters M, Kerkhof W, Kleukers R (2015) De Nederlandse sprinkhanen en krekels (Or- thoptera). Entomologische Tabellen 8, 248 pp. Frequency Additional measures 2-3 months/year Livestock moved to valleys in winter. 1-2 cuts/year Unmown refuges (exclosures). 1 cut/year Uncut refuges (exclosures). Protect from drying. n/a May need to manage scrub encroachment. 1 cut/year Uncut refuges (exclosures). Ungrazed refuges (exclosures). Summer/all year Avoid grazing of very wet areas. 1-2 cuts/year Unmown refuges (exclosures). Beckmann BC, Purse BV, Roy DB, Roy HE, Sutton PG, Thomas CD (2015) Two species with an unusual combination of traits dominate respons- es of British grasshoppers and crickets to environmental change. PLoS ONE 10: e0130488. https://doi.org/10.1371/journal.pone.0130488 Benton T (2012) Grasshoppers and Crickets. Collins, London, 544 pp. Berg H-M, Bieringer G, Zechner L (2005) Rote liste der heuschrecken (Or- thoptera) Osterreichs. In: Zulka K-P (Ed.) Rote Listen gefahrdeter Tiere Osterreichs, BOhlau Verlag, Wien, 167-209. Cheesman O, Brown V (1998) English Nature Species Recovery Pro- gramme - Large Marsh Grasshopper (Stethophyma grossum) 1998 Pre- recovery Project Report. English Nature, Peterborough. JOURNAL OF OrRTHOPTERA RESEARCH 2018, 27(1) 96 Clarke EJ (1948) Studies in the ecology of British grasshoppers. Transac- tions of the Royal Entomological Society of London 99: 173-222. https://doi.org/10.1111/j.1365-2311.1948.tb01235.x Decleer K, Devriese H, Hofmans K, Lock K, Barenbrug B, Maes D (2000) Voorlopige atlas en “rode lijst” van de sprinkhanen en krekels van Belgie (Insecta, Orthoptera). SALTABEL, sprinkhanenwerkgroep van de Benelux i.s.m. Instituut voor Natuurbehoud & Koninklijk Belgisch Instituut voor Natuurwetenschappen, 74 pp. Detzel P (1998) Die Heuschrecken Baden-Wurtembergs. Ulmer, 580 pp. Edwards B (2002) The current status of Stethophyma grossum L. large marsh grasshopper in Dorset. Dorset Environmental Records Centre, Dorchester. English Nature (2005) Grazing management of lowland heathlands. English Nature, Peterborough. http://publications.naturalengland. org.uk/file/ 115020 Evans M, Edmondson R (2007) A Photographic Guide to the Grasshop- pers and Crickets of Britain and Ireland. WGUK, King’s Lynn, 183 pp. Fischer J, Steinlechner D, Zehm A, Poniatowski D, Fartmann T, Beckmann A, Stettmer C (2016) Die Heuschrecken Deutschlands und Nordti- rols. Quelle and Meyer, 368 pp. Fontana P, Kleukers R (2002) The Orthoptera of the Adriatic coast of Italy (Insecta: Orthoptera). Biogeographia — The Journal of Integrative Bio- geography 23: 35-53. https://doi.org/10.21426/B6110182 Galvagni A (2001) Gli Ortotteroidei della Val Venosta, detta anche Vinschgau (Alto Adige, Italia settentrionale). Insecta: Blattaria, Mantodea, Orthop- tera, Dermaptera). Atti Accademia Rovereto Agiati, Rovereto, a. 2001: 251. http://www.agiati.org/UploadDocs/5052_Art04_galvagni_venosta_.pdf Gardiner T (2013) Knee deep to a grasshopper in the New Forest. Country- Side 33: 9-11. http://www.bna-naturalists.org/spring%2013%20]1].pdf Gardiner T, Pye M, Field R, Hill J (2002) The influence of sward height and vegetation composition in determining the habitat preferences of three Chorthippus species (Orthoptera: Acrididae) in Chelmsford, Essex, UK. Journal of Orthoptera Research 11: 207-213. https://doi. org/10.1665/1082-6467(2002)011[0207:TIOSHA]2.0.CO;2 Groome G, Shaw P (2015) Vegetation response to the reintroduction of cattle grazing on an English lowland valley mire and wet heath. Con- servation Evidence 12: 33-39. https://www.conservationevidence. com/teference/download/5501 Grootjans AP, Verbeek SJ (2002) A conceptual model of European wet meadow restoration. Ecological Restoration 20: 6-9. https://doi. org/10.3368/er.20.1.6 Haes E, Harding P (1997) Atlas of Grasshoppers, Crickets and Allied Insects in Britain and Ireland. ITE research publication, no.11, The Stationery Office, London, 61 pp. http://nora.nerc.ac.uk/7358/1/Grasshoppers.pdf Hansen M, Jorgensen O (2010) Insekter i Danmark. Gyldendal, Copenha- gen, 361 pp. Harvey MC, Brock PD (2017) New Forest Large Marsh Grasshopper (Steth- ophyma grossum) 2017 Survey Report. Higher Level Stewardship Agree- ment The Verderers of the New Forest AG00300016. Hochkirch A, Willemse LPM, Rutschmann F, Chobanov DP, Kleukers R, Kristin A, Presa JJ, Szovenyi G (2016) Stethophyma grossum. The IUCN Red List of Threatened Species 2016: e.T16084611A74516611. http:// www.iucnredlist.org/details/summary/16084611/1 Holst K (1986) The Saltatoria (Bush-Crickets, Crickets and Grasshoppers) of Northern Europe. Volume 16 of Fauna Entomologica Scandinavi- ca. Brill, Leiden/Copenhagen, 127 pp. Homburger H, Liischer A, Scherer-Lorenzen M, Schneider MK (2015) Pat- terns of livestock activity on heterogeneous subalpine pastures re- veal distinct responses to spatial autocorrelation, environment and management. Movement Ecology 3: 1-15. https://doi.org/10.1186/ $40462-015-0053-6 Humbert J-Y, Ghazoul J, Walter T (2009) Meadow harvesting techniques and their impact on field fauna. Agriculture, Ecosystems and Environ- ment 130: 1-8. https://doi.org/10.1016/j.agee.2008.11.014 JNCC (2010) UK Priority Species data collation Stethophyma grossum ver- sion 2. http://jncc.defra.gov.uk/_speciespages/588.pdf Kenyeres Z, Szentirmai S (2017) Effects of different mowing regimes on orthopterans of Central European mesic hay meadows. Journal of Or- thoptera Research 26: 29-37. https://doi.org/10.3897/jor.26.14549 J. MILLER AND T. GARDINER Kleukers R, van Nieukerken E, Ode B, Willemse L, van Wingerden W (2004) De Sprinkhanen en Krekels van Nederland (Orthoptera). Nederlandse Fauna 1, 248 pp. Kranebitter P (2008) Die Heuschreckenfauna (Saltatoria, Insecta) des Schlern (Sudtirol). Gredleriana 8: 301-320. https://www.zobodat.at/ pdf/Gredleriana_008_0301-0320.pdf Krause S (1996) Populationsstruktur, Habitatbindung und Mobilitat der Larven von Stethophyma grossum. Articulata 11: 77-89. http://www.dgfo- articulata.de/articulata/1996_2/articulata_1996_11_2_08_krause_.pdf Lake S, Underhill-Day J (2004) Conservation grazing on lowland heaths. The RSPB, Sandy, 95 pp. Lucas WJ (1920) A Monograph of the British Orthoptera. The Ray Society, Adlard and Son and West Newman, London and Dorking, 392 pp. https://doi.org/10.5962/bh1.title.1705 Liiders U-R (2009) Nuevos Datos sobre la distribuci6n de Stethophyma grossum (Linnaeus, 1758) (Orthoptera, Acrididae) en el norte de Es- pana. Boletin Sociedad Entomologica Aragonesa 44: 577-578. http:// sea-entomologia.org/Publicaciones/PDF/BOLN44/577_578BSEA44 NBStethophymagrossum.pdf Maas S, Detzel P, Staudt A (2002) Gefahrdungsanalyse der Heuschrecken Deutschlands. Verbreitungsatlas, Gefahrdungseinstufung und Schutz- konzepte. Bundesamt fur Naturschutz, 402 pp. Malkus J (1997) Habitatpraferenzen und Mobilitat der Sumpfschrecke (Stethophyma grossum L. 1758) unter besonderer Beriicksichtigung der Mahd. Articulata 12: 1-18. http://www.dgfo-articulata.de/articu- lata/1997_1/articulata_1997_12_1_02_malkus_.pdf Marshall J, Haes E (1988) Grasshoppers and Allied Insects of Great Britain and Ireland. Harley Books, Essex, 252 pp. Marzelli M (1997) Untersuchungen zu den Habitatanspruchen der Sumpf- schrecke (Stethophyma grossum) und ihre Bedeutung fur das Habitat management. Articulata 12: 107-121. http://www.dgfo-articulata.de/ articulata/1997_2/articulata_1997_12_2_03_marzelli_.pdf Monnerat C, Thorens P, Walter T, Gonseth Y (2007) Liste rouge des Orthop- teres menacés de Suisse. Office fédéral de l'environnement, Berne et Centre suisse de cartographie de la faune, Neuchatel. Lenvironnement pratique 0719, 62 pp. http://www.unine.ch/files/live/sites/cscf/files/ Documents%20%C3%A0%20t%C3%AI1%C3%AIcharger/Ortho/ liste% 20rouge%20orthopt%C3%A8res.pdf Ortner A, Lechner K (2015) Rote Liste gefahrdeter Heuschrecken Vorarl- bergs. inatura Erlebnis Naturschau - Rote Listen 9, 136 pp. https:// www.zobodat.at/pdf/Rote-Listen-Vorarlbergs_9_0001-0136.pdf Pinchen B, Ward L (2010) The New Forest cicada and other invertebrates. In: Newton A (Ed.) Biodiversity in the New Forest. Pisces Publica- tions, Newbury, 58-64. Ragge D (1965) Grasshoppers, Crickets and Cockroaches of the British Isles. Frederick Warne and Co, London, 300 pp. Sardet E, Roesti C, Braud Y (2015) Cahier d’identification des Orthopteres de France, Belgique, Luxembourg and Suisse. Biotope Editions, 304 pp. Sonneck A, Bonsel A, Matthes J (2008) Der Einfluss von Landnutzung auf die Habitate von Stethophyma grossum an Beispielen aus Mecklenburg- Vorpommern. Articulata 23: 15-30. http://www.dgfo-articulata.de/ articulata/2008_23-1/Articulata_23-1%20Sonneck%20et%20al.pdf Soérens A (1996) Zur Populationsstruktur, Mobilitat und dem Eiablageverh- alten der Sumpfschrecke (Stethophyma grossum) und der Kurzfliigeli- gen Schwertschrecke (Conocephalus dorsalis). Articulata 11: 37-48. Sutton PG (2015) A review of the scarce and threatened Orthoptera and allied species of Great Britain, Orthoptera, Dictyoptera, Dermaptera, Phasmida. Species Status No. 21, Natural England Commissioned Reports, 51 pp. Sutton PG, Beckmann BC, Nelson B (2017) The current status of Orthop- teroid insects in Britain and Ireland. Atropos 59: 6-35. Symes N, Day J (2003) A practical guide to the restoration and manage- ment of lowland heathland. The RSPB, Sandy, 318 pp. Thorens P, Nadig A (1997) Atlas de Distribution des Orthopteres de Suisse. Centre Suisse de Cartographie de la Faune, 236 pp. Voisin J [Coord.] (2003) Atlas des Orthopteres et des Mantides de France. Publications Scientifiques du Museum, 104 pp. Wind P, Pihl S (2010) The Danish Red List. The National Environmental Research Institute. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)