Responses of Plants and Arthropods to Burning and Disking of Riparian Habitats

Abstract: Alteration of Iowa, USA, landscapes for agricultural production has resulted in a loss of >99% of the original prairie and >95% of native wetlands. This conversion has included riparian areas, which, as interfaces between terrestrial and aquatic ecosystems, are important to many wildlife species. Farm Bill programs have resulted in the reestablishment of millions of hectares of grasslands and wetlands nationwide, including >100,000 ha in riparian areas of the Midwest. We assessed plant and arthropod responses to burning and disking of riparian grasslands in east-central Iowa in 2001 and 2002. Burning altered the plant community by removing litter and standing dead vegetation and had negative effects on several arthropod taxa, including Hemiptera and Lepidoptera. However, we observed no differences in vegetation or arthropods between burned and unburned fields during the second year postburning (P > 0.05). Disking decreased the cover of grasses, litter, and standing dead vegetation and increased plant species richness and the cover of forbs and bare ground (P < 0.05). Arthropod abundance and dry biomass were greater on disked than undisked portions of fields (P < 0.05). Increases in the abundance and biomass of arthropods associated with changes in vegetation structure and composition likely improved habitat quality for a number of breeding bird species. Both burning and disking appear to be effective management options for maintaining or enhancing riparian grasslands for wildlife.     

Effects of fire,topography and year-to-year climatic variation on species composition in tallgrass prairie

Native unploughed tallgrass prairie from Konza Prairie, Kansas, USA is described with respect to plant species compositional changes over a five year period in response to fire and topography. The principal gradient of variation in the vegetation is related to time since burning. Species show an individualistic response in terms of relative abundance to this gradient. Both the percentage of and cover of C₄ species and all grasses decrease as the prairie remains unburnt. Forb and woody plant species numbers and abundance increase along this gradient. A secondary gradient of variation reflects topography (i.e. upland versus lowland soils). Upland soils support a higher species richness and diversity. Upland and lowland plant assemblages are distinct except on annually burnt prairie. The interaction between burning regime, topography and year-to-year climatic variation affects the relative abundance of the plant species differentially. The most dominant species overall, Andropogon gerardii, was affected only by year-to-year variation (i.e. climate). Its position at the top of the species abundance hierarchy was unaffected by burning regime or soil type. The other dominant species showed a suite of varying responses to these factors.Deceased May, 1986.     

Restoration of Prairie Community Structure and Ecosystem Function in an Abandoned Hayfield: A Sowing Experiment

Using a multispecies seed sowing experiment, we investigated the roles of seed and microsite limitation in constraining the restoration of native prairie diversity and ecosystem function in an abandoned upland hayfield in northeastern Kansas. Seeds of 32 native and naturalized plant species from the regional pool were sown into undisturbed and experimentally disturbed field plots. After six growing seasons, experimental sowing led to major shifts in species and functional group composition, increases in native species abundance and floristic quality, declines in abundance of non‐native species, and increases in plant diversity. These changes in community structure led to significant changes at the ecosystem level including increases in light capture, peak biomass, primary production, litter biomass, root biomass, and C storage in roots. Our findings reveal the importance of seed limitations in constraining the natural recovery of prairie vegetation, biodiversity, and ecosystem functioning in this grassland and confirm broadcast sowing as a useful tool for the restoration of upland hayfield sites.     

Multi-scale impacts of crested wheatgrass invasion in mixed-grass prairie

Most of North America’s northern Great Plains have been cultivated for crop production, leaving remnants of natural mixed-grass prairie fragmented and threatened by alien plant invasions. The region’s most widespread alien perennial forage crop, crested wheatgrass (Agropyron cristatum sensu amplo), has invaded native grassland and raised concerns regarding its ecological impact. To evaluate impacts at multiple scales of organization, adjacent invaded and uninvaded mixed-grass prairie were sampled at eight widely separated locations. At the population level, native C₃ mid-grasses and forbs were less abundant in invaded grasslands, while native C₃ and C₄ short-grass abundance was not different. At community and landscape levels, diversity was lower in invaded grasslands largely because of lower forb species richness and cover, and crested wheatgrass dominance of both cover (14% basal cover) and seedbank (404 seeds m⁻²). At the ecosystem level, both vegetation and litter biomass were greater in invaded grasslands, however, below ground organic matter (roots and litter), soil organic carbon, total nitrogen and phosphorus were not different. Crested wheatgrass invasion of mixed-grass prairie was associated with lower diversity within and among plant communities, and appears to simplify the composition of mixed-grass prairie landscapes. Hypotheses for crested wheatgrass dominance and persistence following invasion are suggested.     

Effects and mechanism of plant litter on grassland ecosystem: A review

Plant litter is dead, above and below ground; organic material i.e. leaves barks, needles, twigs and roots. Plant litter plays a key role in nutrient cycling and community organization in grassland ecosystems. Litter can have important consequences on recruitment of plant species through modification of biological, physical, and chemical features of microenvironment. Plant litter offers a major input of organic matter to the soil which modifies soil chemistry, hence impacts nutrient cycling. At early stages of litter decomposition, a particular amount of carbon is transporting to the soil nutrient pool. In terrestrial ecosystems, plant litter regulating biogeochemical cycles, maintain soil fertility, nutrient availability, and therefore influence plant growth, diversity, composition, structure, and productivity. Litter can also impact plant above net plant productivity and below net plant productivity in grassland ecosystem. Plant litter accumulation and decomposition can impact plant species composition and community structure through temperature, light and nutrient availability. The effects of plant litter on vegetation may be negative, positive or neutral due vegetation variability, study duration, habitat, latitude, quantity and quality of litter. These diverse effects of plant litter on grassland ecosystem might be due to, management practice type, management intensity, climate type, timing, precipitation and soil nutrient pool etc. Current review attempts to describe prominent effects of plant litter on vegetation, seed germination, soil fertility, Productivity, species composition, community structure and mechanism in grassland ecosystem.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

Long‐term effects of the abandonment of grazing on steppe‐like grasslands

Question: What are the consequences of grazing abandonment on the Stipa lessingiana dominated steppe‐like grasslands? What is the relative importance of management and environmental factors in causing variation in species composition and abundance in the continuously grazed and abandoned grassland stands? Location: Transylvanian Lowland, Romania. Methods: Repeated vegetation mapping of a grassland stand, where grazing was abandoned 35 years ago; re‐sampling six grassland stands surveyed 29–57 years ago. For revealing long‐term changes in species composition and rank abundance PCoA ordination was applied. The relative importance of management and environmental factors in structuring vegetation were explored by CCA ordination. Diversity, evenness and the relative number and abundance of red‐listed species were compared between managed and abandoned stands. Results: Our results pointed out that grasslands which were formerly grazed and dominated by S. lessingiana, in the long‐term absence of grazing, have been transformed into a S. pulcherrima dominated type. Management, probably by creating bare surfaces and preventing litter accumulation, had the strongest effect on the species composition and abundance in the grasslands. Abandoned grassland stands had lower diversity and evenness compared to continuously grazed stands. While at the same time, the relative number of threatened, rare species did not differ between managed and abandoned sites. Conclusion: Maintaining extensively grazed, as well as un‐managed, Stipa dominated grasslands would be important in order to create various habitat conditions for plant species, especially threatened and rare species, and promote diversity on the landscape scale.     

Habitat and landscape effects on abundance of Missouri's grassland birds

Of 6 million ha of prairie that once covered northern and western Missouri, <36,500 ha remain, with planted, managed, and restored grasslands comprising most contemporary grasslands. Most grasslands are used as pasture or hayfields. Native grasses largely have been replaced by fescue (Festuca spp.) on most private lands (almost 7 million ha). Previously cropped fields set aside under the Conservation Reserve Program (CRP) varied from a mix of cool-season grasses and forbs, or mix of native warm-season grasses and forbs, to simple tall-grass monocultures. We used generalized linear mixed models and distance sampling to assess abundance of 8 species of breeding grassland birds on 6 grassland types commonly associated with farm practices in Missouri and located in landscapes managed for grassland-bird conservation. We selected Bird Conservation Areas (BCAs) for their high percentage of grasslands and grassland-bird species, and for <5% forest cover. We used an information-theoretic approach to assess the relationship between bird abundance and 6 grassland types, 3 measures of vegetative structure, and 2 landscape variables (% grassland and edge density within a 1-km radius). We found support for all 3 levels of model parameters, although there was less support for landscape than vegetation structure effects likely because we studied high-percentage-grassland landscapes (BCAs). Henslow's sparrow (Ammodramus henslowii) counts increased with greater percentage of grassland, vegetation height-density, litter depth, and shrub cover and lower edge density. Henslow's sparrow counts were greatest in hayed native prairie. Dickcissel (Spiza americana) counts increased with greater vegetation height-density and were greatest in planted CRP grasslands. Grasshopper sparrow (A. savannarum) counts increased with lower vegetation height, litter depth, and shrub cover. Based on distance modeling, breeding densities of Henslow's sparrow, dickcissel, and grasshopper sparrow in the 6 grassland types ranged 0.9–2.6, 1.4–3.2, and 0.1–1.5 birds/ha, respectively. We suggest different grassland types and structures (vegetation height, litter depth, shrub cover) are needed to support priority grassland-bird species in Missouri. © 2011 The Wildlife Society.     

Micro-Scale Restoration: A 25-Year History of a Southern Illinois Barrens

We studied vegetation change of a remnant barrens in southern Illinois over twenty‐five years. The study area was periodically burned between 1969 and 1993, but fire was excluded for a 16‐year period (1974–1989). During the study, the barrens supported a mixture of species whose preferred habitats ranged from prairie and open woodlands to closed forest communities. The herbaceous vegetation may be on a trajectory characterized by increasing dominance of woodland species and declining prairie species. Fire management temporarily reversed this trend, but it continued once fire was excluded. Reintroduction of prescribed burning in 1990–1993 altered the vegetation trajectory but not back toward a species composition comparable to that present on the site before cessation of fire management after 1973. Following interruption of prescribed burning, tree basal area more than doubled, and density showed a 67% increase between premanagement conditions in 1968 and 1988. Salix humilis (prairie willow) density had significant negative correlations with tree density and basal area. However, there was no consistency in response of shrub species on the site to the varied site conditions over time. Fire management on the site may not recover the historic barrens that occurred on the site. Nevertheless, consistent fire management will drive vegetation changes toward increasing abundance of prairie and open woodland species that would otherwise be lost without burning.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.     

Frequent fire promotes diversity and cover of biological soil crusts in a derived temperate grassland

The intermediate disturbance hypothesis (IDH) predicts that species diversity is maximized at moderate disturbance levels. This model is often applied to grassy ecosystems, where disturbance can be important for maintaining vascular plant composition and diversity. However, effects of disturbance type and frequency on cover and diversity of non-vascular plants comprising biological soil crusts are poorly known, despite their potentially important role in ecosystem function. We established replicated disturbance regimes of different type (fire vs. mowing) and frequency (2, 4, 8 yearly and unburnt) in a high-quality, representative Themeda australis–Poa sieberiana derived grassland in south-eastern Australia. Effects on soil crust bryophytes and lichens (hereafter cryptogams) were measured after 12 years. Consistent with expectations under IDH, cryptogam richness and abundance declined under no disturbance, likely due to competitive exclusion by vascular plants as well as high soil turnover by soil invertebrates beneath thick grass. Disturbance type was also significant, with burning enhancing richness and abundance more than mowing. Contrary to expectations, however, cryptogam richness increased most dramatically under our most frequent and recent (2 year) burning regime, even when changes in abundance were accounted for by rarefaction analysis. Thus, from the perspective of cryptogams, 2-year burning was not an adequately severe disturbance regime to reduce diversity, highlighting the difficulty associated with expression of disturbance gradients in the application of IDH. Indeed, significant correlations with grassland structure suggest that cryptogam abundance and diversity in this relatively mesic (600 mm annual rainfall) grassland is maximised by frequent fires that reduce vegetation and litter cover, providing light, open areas and stable soil surfaces for colonisation. This contrasts with detrimental effects of 2-year burning on native perennial grasses, indicating that this proliferation of cryptogams has potentially high functional significance for situations where vegetation cover is depleted, particularly for reducing soil erosion.     

Old‐field grassland successional dynamics following cessation of chronic disturbance

Question: Does increasing Festuca canopy cover reduce plant species richness and, therefore, alter plant community composition and the relationship of litter to species richness in old‐field grassland? Location: Southeastern Oklahoma, USA. Methods: Canopy cover by species, species richness, and litter mass were collected within an old‐field grassland site on 16, 40 m × 40 m plots. Our study was conducted during the first three years of a long‐term study that investigated the effects of low‐level nitrogen enrichment and small mammal herbivory manipulations. Results: Succession was altered by an increase in abundance of Festuca over the 3‐yr study period. Species richness did not decline with litter accumulation. Instead, Festuca increased most on species‐poor plots, and Festuca abundance remained low on species‐rich plots. Conclusions: Festuca may act as an invasive transformer‐species in warm‐season dominated old‐field grasslands, a phenomenon associated more with invasions of cool‐season grasses at higher latitudes in North America.     

Site‐specific responses of native and exotic species to disturbances in a mesic grassland community

Abstract. Grassland communities are increasingly recognized as disturbance‐dependent ecosystems, yet there are few replicated, multi‐site studies documenting vegetation responses to varying frequencies and types of grassland disturbance. Even so, land managers frequently manipulate disturbance regimes in an attempt to favour native grassland plants over exotic species. We conducted a factorial experiment testing three frequencies of clipping combined with litter accumulation, litter removal, and soil disturbance within the highly threatened California coastal prairie plant community. We monitored the response of native/exotic, grass/forb plant guilds once a year for four years. More frequent clipping reduced cover of exotic grasses and favoured exotic forbs, whereas native species were largely unaffected by clipping frequency. Litter accumulation, litter removal, and soil disturbance did not affect vegetation composition. Effects of litter accumulation may take longer than our experiment allowed, and soil disturbance due to our treatments was not sufficiently strong to show consistent effects relative to mammalian soil disturbance. Treatment response of some plant guilds differed among sites, highlighting the importance of replicating experiments at several sites before recommending conservation management practices.     

Soil physiochemical changes following 12 years of annual burning in a humid–subtropical tallgrass prairie: a hypothesis

Burning is known to stimulate growth of grassland vegetation, promote species diversity, and inhibit natural invasion by woody plants. However, the frequency at which grasslands are burned as part of their management can affect soil nutrient content and, ultimately, productivity. The objective of this study was to characterize changes in soil physical and chemical properties in a native tallgrass prairie after 12 years of annual burning. In 1989, five soil samples from the 0 to 10 cm depth were collected along a transect through a 3 ha parcel of native tallgrass prairie in central Arkansas. Soil sampling was repeated in 2001 to assess changes over time. Results showed that soil bulk density, electrical conductivity, extractable P, Na, Fe, and Mn decreased significantly (P < 0.05), while soil organic matter, total N and C, and the C/N ratio increased significantly (P < 0.05) within the 12-year period during which annual burning was the only imposed management practice. Mean extractable K, Ca, Mg, S, and Zn levels were all lower in 2001 than in 1989, but differences were not significant, while soil pH did not change. The results of this study indicate that annual export of several essential plant nutrients during prescribed burning of relatively small, remnant prairie fragments exceeds annual imports from atmospheric deposition and/or organic matter mineralization. Annual prescribed burning may be too frequent to maintain optimal ecosystem functioning and productivity. Decreasing the frequency of prescribed burning for native grassland management may help to retain more soil nutrients to sustain a higher level of productivity.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.     

Prairie vegetation and soil nutrient responses to ungulate carcasses

The impact of large ungulate carcasses on grassland dynamics was investigated by monitoring vegetation and soil nutrients in 50-cm circular zones around the center of bison (Bos bison), cattle (B. taurus), and deer (Odocoileus virginianus) carcasses. An ungulate carcass creates an intense localized disturbance that varies with animal size and the season of death. Unlike other natural disturbances, carcasses deposit a concentrated pulse of nutrients into the soil. One year after death, inorganic nitrogen concentrations were significantly higher in the inner 50 cm at both adult and juvenile carcass sites than in surrounding prairie. Areas around a carcass became zones of fertility that favored different components of the vegetation and stimulated biomass production. Species richness and diversity at the center of carcass sites were lowest 1 year after death, but increased significantly in subsequent years. However, warm-season perennial grasses declined near the center of carcass sites and did not recover. Five years after death, ungulate carcass sites remained disturbed patches that harbored vegetation characteristically different in composition and stature from surrounding prairie. By providing a niche for species not normally found in undisturbed prairie, carcasses increased community heterogeneity and may play an important role in adding spatial complexity to grassland ecosytems. Received: 12 April 1999 / Accepted: 7 September 1999     

Invertebrate,not small vertebrate,herbivory interacts with nutrient availability to impact tallgrass prairie community composition and forb biomass

The effects of herbivores and their interactions with nutrient availability on primary production and plant community composition in grassland systems is expected to vary with herbivore type. We examined the effects of invertebrate and small vertebrate herbivores and their interactions with nutrient availability on grassland plant community composition and aboveground biomass in a tallgrass prairie ecosystem. The abundance of forbs relative to grasses increased with invertebrate herbivore removals. This increase in forb abundance led to a shift in community composition, where invertebrate removals resulted in greater plant species evenness as well as a divergence in composition among plots. In contrast, vertebrate herbivore removals did not affect plant community composition or aboveground biomass. Nutrient additions alone resulted in a decrease in plant species richness and an increase in the abundance of the dominant grass, but the dominant grass species did not greatly increase in abundance when nutrient additions were combined with invertebrate removals. Rather, several subdominant forbs came to dominate the plant community. Additionally, the combined nutrient addition and invertebrate herbivore removal treatment increased forb biomass, suggesting that invertebrate herbivores suppress the responses of forb species to chronic nutrient additions. Overall, the release of forbs from invertebrate herbivore pressure may result in large shifts in species composition, with consequences for aboveground biomass and forage quality due to altered grass:forb ratios in grassland systems.     

Long-Term Grazing Accelerated Litter Decomposition in Northern Temperate Grasslands

Livestock grazing affects plant community composition, diversity, and carbon (C) and nutrient cycling in grasslands. Grazing leads to plant communities that have higher relative abundance of grazing-tolerant species, which in turn may alter the chemical composition of biomass and subsequent litter decomposition rates. To better understand the effects of long-term grazing and associated vegetation shifts on biogeochemical cycling in northern temperate grasslands of western Canada, we studied litter decomposition over 18 months at 15 locations, stratified across the Mixed-grass Prairie, Central Parkland, and Foothills Fescue natural subregions. At each location, we examined decomposition in an area exposed to grazing and an area where cattle were excluded. We used litterbags containing leaf litter from seven major grass species representing different grazing tolerances and included a local source of community litter from each study site and cellulose paper as standards. Decomposition was affected by litter types, with litter from grazing-tolerant species such as Poa pratensis and Bouteloua gracilis having faster decomposition rates compared to grazing-intolerant species, supporting the hypothesis that changes in vegetation composition due to grazing influences biogeochemical cycling by modifying litter decomposition in grasslands. Litter decomposition was also overall most rapid in the cool–wet Foothills Fescue, followed by the temperate mesic Central Parkland, and slowest in the warmer–drier Mixed-grass Prairie. Combined with known grazing-induced changes in grassland composition, these findings indicate that livestock grazing may accelerate litter decomposition rates in the more mesic Foothills Fescue and parkland regions, but not the more arid Mixed-grass Prairie. Overall, this study elucidates the role of livestock grazing and its associated effects on litter decomposition and ecosystem processes in northern grassland ecosystems.     

Effects of the litter layer of Pteridium aquilinum on seed banks under experimental restoration

Questions: Does the litter layer of Pteridium aquilinum (bracken) act as a barrier to certain species in the seed bank? Does bracken control/restoration treatment affect seed transfer through the litter layer? Location: Five experiments at three sites across the UK covering two major vegetation types; acid‐grassland and heath‐land. Methods: At each experiment a range of bracken control and vegetation restoration treatments were applied for about ten years. The seed bank was sampled in both the bracken litter and the soil. The cover (%) of each species in the vegetation and the bracken litter abundance (cover and depth) was also estimated. Results: The bracken litter layer acts as an inert barrier as it contained a large proportion of seeds available in the litter‐soil profile (38%– 67% of the total). Bracken litter depth and cover also influenced significantly the seed bank composition in both the bracken litter and the soil. These effects were site‐specific, and species‐specific. The application of treatments changed significantly the balance between seed inputs and outputs in the bracken litter layer for some species. This was either a positive or negative response relative to the untreated control plots. Conclusion: For heathland and acid‐grassland restoration, the bracken litter layer may be an important seed source, but it must be disturbed particularly before seed addition.