Maintain species-rich, semi-natural grassland
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Overall effectiveness category Awaiting assessment
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Number of studies: 19
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Supporting evidence from individual studies
A replicated, paired, site comparison study in 1990–1991 in 35 calcareous grasslands in Northern Bavaria, Germany (Völkl et al. 1993) reported that semi-natural grasslands maintained by sheep grazing had a higher density of meadow neb moth Metzneria metzneriella caterpillars, and a similar occurrence of hoary bell moth Eucosma cana caterpillars, compared to mown grasslands. Results were not tested for statistical significance. In grazed grasslands, 2.9–3.3% of greater knapweed Centaurea scabiosa flowerheads contained meadow neb caterpillars, compared to 0–0.3% of flowerheads in mown grasslands, and 2.2–2.5% of flowerheads in abandoned grasslands. The occurrence of hoary bell was similar in mown, grazed and abandoned grasslands (data not presented). Thirty-five grasslands (0.5–2 ha) were managed by either light sheep grazing in early autumn (7 sites, vegetation <10 cm) or annual mowing (usually in midsummer, 7 sites, vegetation ~25 cm before cutting), or had been abandoned for at least five years (21 sites, vegetation >25 cm with shrubs). In September–October 1990 and 1991, samples of 100–350 greater knapweed flowerheads/site were collected from seven pairs of grazed-abandoned and mown-abandoned grasslands, and seven (1990) and four (1991) unpaired, abandoned grasslands. Flowerheads were dissected in the laboratory to identify caterpillars.
Study and other actions testedA replicated, site comparison study in 1994 in 19 traditional hay meadows in Bavaria, Germany (Dolek & Geyer 1997) found that the abundance and species richness of all butterflies, and of threatened species only, was similar in mown and grazed grassland. In lightly grazed meadows, the abundance of all species of butterfly (39.5 individuals) and of 16 threatened species (6.9 individuals) was similar to the abundance in meadows mown once/year (all species: 25.8 individuals; threatened species: 7.5 individuals). The species richness of butterflies was also similar in grazed and mown meadows (data not presented). However, managed meadows had a higher abundance and species richness of butterflies than abandoned meadows (data not presented). Nineteen meadows, which had been managed in the same way for at least 5–20 years, were compared. Six traditionally managed hay meadows were mown once/year in July or early August, nine meadows were extensively grazed with sheep, cattle or horses for a few weeks each summer, one meadow was grazed by sheep throughout the summer, and three meadows were not managed (abandoned). From June–August 1994, butterflies were surveyed along a fixed transect five times in each meadow.
Study and other actions testedA replicated, site comparison study in 1993–1994 in 16 alpine meadows in southern Switzerland (Schwarzwälder et al. 1997) found that traditional hay meadows and recently abandoned meadows had a higher abundance of heath fritillary Mellicta athalia adult males and caterpillars, but not females, than old, abandoned or restored meadows. There were more adult males and caterpillars in traditional hay meadows (males peak: 30 individuals/hour; caterpillars: 0.5–3.5 individuals/hour) and recently abandoned meadows (males: 40 individuals/hour; caterpillars: 4–8 individuals/hour) than in old, abandoned (males: 21 individuals/hour; no caterpillars) or restored meadows (males: 20–22 individuals/hour; caterpillars: 0–0.2 individuals/hour). The number of females was not significantly different between meadows (traditional: 5; recently abandoned: 14; old abandoned: 5; restored: 7–14 individuals/hour). Marked butterflies were recorded moving between all habitat types. Five traditional hay meadows were mown once/year in June or July, five old, abandoned meadows had been unmanaged for >25 years, two recently abandoned meadows had been unmanaged for around six years. From 1992, two abandoned meadows were restored by annual mowing, and two were restored by mowing every 4–5 years. From June–July 1993–1994, adult butterflies were caught and marked for 45 minutes/meadow every other day. In 1994, each meadow was searched for three hours, spread over several days, to record solitary caterpillars.
Study and other actions testedA replicated, site comparison study in 1992 in agricultural land in Baden-Württemberg, Germany (Steffan-Dewenter & Tscharntke 1997) found that old meadows had a higher species richness of adult butterflies and caterpillars than either recently established set-aside. Adult butterfly species richness was higher in old meadows (20 species) than in naturally regenerated set-aside (11–13 species), sown set-aside (7 species) or cereal crops (4 species). Caterpillar species richness was also higher in old meadows (16 species) than in naturally regenerated set-aside (3–7 species). Butterfly species found in meadows tended to be less migratory, spend longer as caterpillars (meadow: 121 days; set-aside: 44–105 days), and have fewer generations/year (meadow: 1.8 generations/year; set-aside: 1.9–2.7 generations/year) than species in recently established set-aside. In 1992, four fields in each of seven management types were studied: old meadows (>30 years old), former cereal fields left to naturally develop as set-aside for each of 1, 2, 3 and 4 years, 1-year-old set-aside sown with lacy phacelia Phacelia tanacetifolia, and cereal fields (rye Secale cereale or wheat Triticum aestivum). Meadows and set-aside fields were mown once/year in July. From May–October 1992, adult butterflies were counted along transects nine times/field. In September 1992, moth and butterfly caterpillars were sampled twice by sweep-netting.
Study and other actions testedA replicated, site comparison study in 1988–1996 in 17 upland prairies in Missouri, Minnesota, North Dakota and Wisconsin, USA (Swengel & Swengel 1997, same experimental set-up as Swengel 1998 and Swengel & Swengel 2001) found that prairies managed by haying or grazing had a higher abundance of four specialist butterfly species, but a lower abundance of three species than prairies managed by burning or unmanaged sites. Of seven prairie specialist butterfly species, three (regal fritillary Speyeria idalia, Pawnee skipper Hesperia leonardus pawnee, Dakota skipper Hesperia dacotae) were more abundant in prairies managed by haying than in rotationally burned, grazed or unmanaged prairies in at least one of three regions. Gorgone checkerspot Chlosyne gorgone was more abundant in grazed prairies than burned or unmanaged areas. However, three species (gray copper Lycaena dione, arogos skipper Atrytone arogos, Poweshiek skipperling Oarisma poweshiek) were less abundant in hayed or grazed prairies than in unmanaged prairies in at least one of three regions. See paper for individual species data. Across 17 prairies (16 to >120 ha), two areas were managed by grazing, six by haying (often in rotation), eight by burning on rotation, three by burning and haying, and two were unmanaged. From 1988–1996, butterflies were surveyed on transects through different management areas at each site. Sites were not surveyed in every year.
Study and other actions testedA replicated, site comparison study in 1986–1995 in 104 tallgrass prairies and 141 pine barrens in Illinois, Iowa, Minnesota, Missouri, North Dakota and Wisconsin, USA (Swengel 1998, same experimental set-up as Swengel & Swengel 1997 and 2001) found that hayed, mown, cut or grazed grasslands had a higher abundance of six of 16 specialist butterfly species, but a lower abundance of three specialist species, than burned or unmanaged grasslands. Of 16 prairie or pine barren specialist butterfly species, five were more abundant in sites managed by haying (Dakota skipper Hesperia dacotae, pawnee skipper Hesperia leonardus pawnee), mowing (Persius duskywing Erynnis persius), cutting (cobweb skipper Hesperia metea) or grazing and haying (regal fritillary Speyeria idalia), than burned or unmanaged sites. Arogos skipper Atrytone arogos was most abundant in grazed and hayed, or unmanaged, sites. Poweshiek skipper Oarisma poweshiek was less abundant in sites managed by haying than in unmanaged sites, and Ottoe skipper Hesperia ottoe and dusted skipper Atrytonopsis hianna were less abundant in sites managed by grazing or mowing than in rotationally burned sites. See paper for individual species data. Seven species had similar abundance between management types (see paper for details). Of 104 prairies (1–2,024 ha), 27 were hayed in summer on a 1–2-year rotation, of which two were also grazed occasionally with cattle; 10 were grazed; 61 were managed by cool-season burning on a 2–5-year rotation, of which 21 were additionally mown or hayed; and six had not been managed for many years. Of 141 pine barrens, some were burned by wildfires, some were used for off-road vehicle trails, and some were power line rights-of-way (no further detail provided). From April–September 1986–1995, butterflies were surveyed on transects at each site. Most sites were surveyed more than once/year, and in >1 year.
Study and other actions testedA replicated, site comparison study in 1990–1997 in 106 tallgrass prairies in Illinois, Iowa, Minnesota, Missouri, North Dakota and Wisconsin, USA (Swengel & Swengel 2001, same experimental set-up as Swengel & Swengel 1997 and Swengel 1998) found that in some states, prairies managed by haying had the highest abundance and species richness of specialist and grassland butterflies, but in other states grazing supported a higher abundance and species richness of butterflies compared to other management types. In Missouri, the abundance and species richness of specialist and grassland butterflies was higher in prairies managed by two-year rotational haying than in rotationally burned areas. In Minnesota, North Dakota and western Iowa, the abundance of specialist and grassland species was higher in hayed areas than in burned areas, and the abundance and richness of specialist species was lower in rotationally grazed areas. However, in eastern Iowa, Illinois, and Wisconsin, the abundance of specialist and grassland species, and the richness of specialists, was higher at a continuously grazed site than in other management types. All data were presented as models results. Across all prairies, specialist and grassland butterfly abundance and richness tended to be higher at rotationally managed sites (haying, grazing or burning) longer after they were last managed. Of 106 prairies (1.2–2,024 ha), 27 contained areas managed by haying, mostly on a two-year rotation, seven were managed by rotational grazing (0.3–0.6 animals/ha/year) and one by continuous grazing (3–6 animals/ha/year), 77 areas were managed by rotational burning (every 2–5 years) in the cool-season (of which 24 were also hayed or mown), and nine had been unmanaged for many years. From May–September 1990–1997, butterflies were surveyed on parallel transects (5–10 m apart) at each site. Most sites were surveyed more than once/year, and in >1 year. Species were classified as “specialists” (of native plants), “grassland” (occurring widely in open habitat) and “generalist” (occurring in a range of habitats).
Study and other actions testedA replicated, site comparison study in 1997–1999 in 16 pastures and meadows in northwest Russia and southeast Finland (Saarinen & Jantunen 2005) found that butterfly abundance, species richness and diversity were similar in mown meadows and grazed pastures. In mown meadows, the total abundance (3,660 individuals) and species richness (46 species) of butterflies was not significantly different from the total abundance (2,082 individuals) and species richness (42 species) in grazed pastures (see paper for diversity data). Only three out of 37 species showed a significant preference for mown meadows (Amanda’s blue Polyommatus amandus, large skipper Ochlodes sylvanus and ringlet Aphantopus hyperantus, see paper for data). The remaining 34 species did not show a significant preference for either field type. Butterfly communities were affected more by the origin and age of the grassland than the present management method (see paper for details). Eight meadows were mown annually in late July or August and eight pastures were grazed by cattle, although some had sheep or horses temporarily. Tilling and fertilisation (manure) tended to occur at intervals of 3–10 years. In June–July 1997–1999, butterflies were surveyed 12–13 times/site along transects (640–720 m).
Study and other actions testedA replicated, site comparison study in 2003 in 38 meadows in Hebei Province, China (Liu et al. 2006) found that uncultivated, grazed meadows were less likely to be occupied by marsh fritillary Euphydryas aurinia eggs and caterpillars than lightly cultivated meadows with grass margins and intercrop, but caterpillar survival was higher in the uncultivated meadows. Fewer entirely uncultivated, grazed meadows contained eggs (1/12 meadows) and caterpillars (5/22 meadows) than meadows with some cultivation (eggs: 9/11; caterpillars: 11/16 meadows). In total, 70 egg clusters were found in grazed meadows, compared to 179 egg clusters in cultivated meadows (statistical significance not assessed). The mortality of egg clusters was similar in grazed meadows (16% of 69 clusters) and cultivated meadows (10% of 177 clusters), but the survival of pre-hibernation caterpillars was higher in grazed meadows (21/59, 33%) than in cultivated meadows (23/164, 14%). A total of 38 meadows (0.025 ha–3.200 ha) were studied. In 2003, twenty-two meadows were entirely uncultivated and grazed. Another 16 meadows contained some cultivation (corn or potatoes), and were divided into cultivated habitat (grass strips within and around the crop, no grazing from April–October) and meadow habitat (meadows and fallow land, grazed by sheep and cattle). In June 2003, twelve uncultivated and 11 cultivated meadows were searched for egg clusters. These were marked and observed every other day until all hatched caterpillars had disappeared or begun overwintering. In September 2003, all 38 meadows were surveyed for caterpillar nests.
Study and other actions testedA replicated, site comparison study in 2004–2005 in two remnant prairies and adjacent land in Iowa, USA (Vogel et al. 2007) found that prairies which were grazed and burned had a higher abundance, but not species richness, of butterflies than prairies which only received grazing or burning, but the three management practices supported different species. The abundance of butterflies in grazed and burned prairies (31.5 individuals/unit) was higher than in prairies which were only grazed (27.8 individuals/unit) or only burned (20.2 individuals/unit). Species richness of butterflies was similar in prairies managed by grazing and burning (8.5 species/unit), only grazing (7.4 species/unit) and only burning (8.6 species/unit). Butterfly diversity was lower in prairies managed by grazing only, or grazing and burning, than in prairies managed by burning only (data presented as model results). However, each management practice supported different species (see paper for details). Across two remnant prairie reserves (320 and 1,800 ha) and surrounding land, 28 management units (10–167 ha) were managed consistently for ≥4 years. Six units were lightly grazed on rotation (1 cow-calf pair/4 ha). Ten units were burned during autumn or spring every 1–6 years. Twelve units were burned and grazed. From June–August 2004–2005, butterflies were surveyed for 30 minutes twice/year at 69 sites (50 × 50 m, >150 m apart) across the 28 units.
Study and other actions testedA site comparison study in 2008–2009 on two semi-natural grasslands in southern Italy (D’Aniello et al. 2011) found that a grassland grazed with sheep and cattle had a higher abundance and species richness of butterflies than a grassland cut for hay. The grazed grassland had a higher abundance (6,005 individuals) and species richness (45 species) of butterflies than the mown grassland (abundance: 2,416 individuals; richness: 28 species). All species found in the mown grassland were also found in the grazed grassland, and most species were more abundant at the grazed site (23/28). See paper for data on individual species. Two 6-ha grasslands surrounded by woodland, both at 850 m altitude and 3 km apart, were studied. One site was mown for hay once/year, in June, and the other site was grazed with sheep and cattle. Both meadows had received the same management for at least 20 years. From April–September 2008–2009, butterflies were surveyed on a weekly 1-km transect around the edge of each site.
Study and other actions testedA replicated, site comparison study in 2003–2005 in 47 alpine meadows in Picos de Europa, Spain (Dover et al. 2011) found that mown hay meadows had a higher abundance of two out of 44 butterfly species than summer grazed or abandoned meadows, and summer grazed meadows had a lower abundance or occurrence of four out of 44 butterfly species than mown or abandoned meadows. The abundance of two species (black-veined white Aporia crataegi and meadow brown Maniola jurtina) was higher in hay meadows than in grazed or abandoned meadows (data presented as model results). The abundance of three species (small skipper Thymelicus sylvestris, small white Pieris rapae, ringlet Aphantopus hyperantus) was lower, and one species (small pearl-bordered fritillary Boloria selene) occurred less frequently, in summer grazed meadows than in hay meadows or abandoned meadows (data presented as model results). Two species (grizzled skipper Pyrgus malvae and painted lady Vanessa cardui) occurred more frequently in managed meadows than in abandoned meadows (data presented as model results). The remaining 36 species did not differ in abundance or occurrence between management types. From summer 2003–2005, management was recorded on 47 meadows. Seven meadows were grazed by livestock in summer, 24 meadows were cut for hay, and 16 meadows were either abandoned or only grazed in the winter. The abandoned meadows had different amounts of scrub growing within them. From June–July 2004, butterflies were surveyed nine times on a transect around the edge of each meadow.
Study and other actions testedA replicated, site comparison study in 2008 in a lowland agricultural landscape in Kanton Fribourg, Switzerland (Haaland & Bersier 2011) found that extensively managed meadows had a lower abundance of butterflies, but a similar species richness and a different community composition, compared to sown wildflower strips. The abundance of butterflies in extensively managed meadows (0.12 individuals/m) was lower than in sown wildflower strips (0.29 individuals/m), but the species richness was similar in meadows (0.05 species/m) and wildflower strips (0.07 species/m). The species composition was different between the two habitats, with six of 25 species observed most frequently in the meadows and seven species occurring only in wildflower strips. None of the five rarest species in the region were recorded in either the meadows or wildflower strips. See paper for details on individual species. Eleven meadows (0.21–1.64 ha) were cut at least twice/year after mid-June. Twenty-five wildflower strips (0.15–1.16 ha) were sown with a standard seed mixture of 24 plant species, and were 1–7 years old. From May–September 2008, butterflies were surveyed once/month on a transect through the middle of each meadow (85–310 m) or wildflower strip (70–450 m).
Study and other actions testedA replicated, site comparison study in 2008 in a mixed farming region in Hungary (Kovács-Hostyánszki et al. 2011) found that established semi-natural grassland supported a higher abundance and species richness of butterflies than cereal fields. In semi-natural grassland fields both the abundance (74 individuals/field) and species richness (8 species/field) of butterflies were higher than in winter wheat fields (abundance: 4 individuals/field; richness: 2 species/field). See paper for details of individual species. From at least 2005, six established semi-natural grassland fields had no fertilizer or chemicals applied, and were either lightly grazed or mown once/year in May–June. Sixteen winter wheat fields were fertilized (70 kg/ha/year nitrogen), sprayed once/year in spring with herbicide and insecticide, and harvested in June. From May–August 2008, butterflies were surveyed on fixed transects four times in each field. Each field was surveyed for 10, 20 or 30 minutes, depending on field size.
Study and other actions testedA replicated, site comparison study in 2007 in a grassland and woodland reserve in the Czech Republic (Slamova et al. 2013) found that grasslands managed by occasional mowing had a similar abundance of Scotch argus Erebia aethiops to grasslands managed by sheep and goat grazing. On occasionally mown grasslands, the abundance of Scotch argus males (9 individuals/ha) and females (5 individuals/ha) was similar to grazed grasslands (males: 7; females: 4 individuals/ha). However, the abundance of Scotch argus males (19 individuals/ha) and females (13 individuals/ha) was highest on temporarily abandoned grasslands, and lowest on intensively mown grasslands (males: 3; females: 2 individuals/ha). Within a 55-ha reserve, 27 grasslands (128–6,072 m2) were managed by either occasional mowing, sheep and goat grazing or intensive mowing, or were temporarily abandoned. On 33 days from July–August 2007, butterflies were caught, individually marked, and recaptured at each site.
Study and other actions testedA replicated, site comparison study in 2007–2010 in 28 grassland sites in Bílé Karpaty Protected Landscape Area, Czech Republic (Bonari et al 2017) found that the management of semi-natural grassland affected the species richness and composition of butterfly and moth communities differently. The species richness of moths was highest in mown grasslands and lowest in grazed grasslands, and these sites had different species composition. However, the species richness of butterflies was highest under mixed management, though species composition was not affected (all data presented as model results). One of four different management practices (mown once/year; grazed by sheep, cattle or deer; abandoned (no grazing or mowing); or ‘mixed’ management) was applied to each of 28 sites (1.5–70.7 ha) for at least five consecutive years. ‘Mixed’ management included mowing different parts of the site at different times, often with patches left uncut for a year, or mowing followed by grazing. From 2007–2010, butterflies and moths were surveyed on >6 visits between April and October in each of two consecutive years to each site.
Study and other actions testedA replicated, site comparison study in 2015 in 20 grasslands in Saxony, Germany (Ernst et al. 2017) found that managed grasslands had a higher abundance of farmland butterflies and burnet moths, but a lower abundance of woodland butterflies and burnet moths, than abandoned grasslands, but there was no difference in species richness or community composition. In managed grasslands, the abundance of 35 species of farmland butterflies and burnet moths (195–206 individuals) was higher than in abandoned grasslands (127 individuals). However, the abundance of 20 species of woodland butterflies and burnet moths was lower in managed grasslands (17–36 individuals) than abandoned grasslands (34–44 individuals). The species richness of both farmland and woodland species was similar in managed (farmland: 14; woodland: 5–6 species) and abandoned (farmland: 13–14; woodland: 5–6 species) grassland. The community composition was also similar in managed and abandoned grasslands (data presented as model results). Twenty calcareous grasslands (0.90–5.38 ha) were surveyed. Eight were managed by summer grazing (May–September, <1 animal/ha, with cattle, sheep, goats, horses or donkeys), one was managed by mowing, one was mown and grazed, and 10 were abandoned (not grazed or mown). From May–August 2015, butterflies and burnet moths were surveyed three times along a 20-minute transect on a 0.8 ha patch at each site. Butterflies and burnet moths were classified as 35 farmland and 20 woodland species.
Study and other actions testedA replicated, site comparison study in 2013–2015 in 45 semi-natural grasslands in eastern Austria (Fiedler et al. 2017) found that grasslands managed by grazing, early mowing and late mowing had distinct butterfly and day-flying moth communities. Butterfly and day-flying moth communities in semi-natural grasslands managed by extensive grazing were different to communities in early-mown and late-mown meadows (data presented as model results). In addition, some species showed a preference for sites that were grazed (crepuscular burnet Zygaena carniolica, transparent burnet Zygaena purpuralis/minos), early-mown (marbled white Melanargia galathea, meadow brown Maniola jurtina) or late-mown (short-tailed blue Cupido argiades). The use of all three grassland management regimes (grazing, early mowing and late mowing) in different parts of the landscape increased butterfly diversity across the landscape (data presented as model results). Semi-natural grasslands managed in three ways were studied: extensive pastures grazed by cattle from April–October, meadows mown once/year in early summer with cuttings removed, and former vineyards mown once/year in late summer with cuttings not removed. In June 2013–2015, all butterflies, burnet moths (Zygaenidae) and hummingbird hawk-moths Macroglossum stellatarum were counted once on 9–11 sites/year (50 × 50 m) under each management type.
Study and other actions testedA replicated, site comparison study in 2014 in 26 grasslands in Germany (Mangels et al. 2017) found that grasslands managed by mowing (sometimes alongside grazing) had a lower abundance, species richness and diversity of moths than unmown, grazed grasslands, but grasslands managed by grazing (sometimes alongside mowing) had a similar abundance, species richness and diversity of moths to ungrazed, mown grasslands. Mown grasslands had a lower abundance, species richness and diversity of moths, and more generalist and widespread species, than unmown, grazed grasslands (data presented as model results). Grazed grasslands had a similar abundance, species richness and diversity of moths to ungrazed, mown grasslands (data presented as model results). In addition, grazed and mown grasslands were inhabited by different moth communities (see paper for details). Of 87 individual species monitored, 10 species preferred mown grasslands and 19 species avoided mown grasslands, while 12 species preferred grazed grasslands and 24 species avoided grazed grasslands (see paper for individual species data). From 2006, across three regions, nine grasslands were managed by grazing (by cattle, sheep or horses at 26–520 livestock units/ha/year), nine by mowing (1–2 cuts/year, often with nitrogen fertilization), and eight were grazed and mown (76–163 livestock units/ha/year; 1–2 cuts/year). Moths were collected once/month from nine grasslands in each of two regions (May–August 2014), and from eight grasslands in one region (June–July 2014). Each night, a 12 V actinic and black-light trap were placed in the centre of each of three grasslands for 138–317 minutes/night.
Study and other actions tested
Where has this evidence come from?
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This Action forms part of the Action Synopsis:
Butterfly and Moth ConservationRelated Actions
| Actions | Effectiveness | Studies | Category | |
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Maintain species-rich, semi-natural grassland Action Link |
Awaiting assessment | 19 |
Synopsis Link |
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Cease grazing on grassland to allow early succession Action Link |
Evidence not assessed | 26 |
Synopsis Link |
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Increase grazing intensity or cutting frequency on grassland Action Link |
Evidence not assessed | 5 |
Synopsis Link |
Butterfly and Moth Conservation - Published 2023
Butterfly and Moth Synopsis
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