Intact Grassland Research Articles

Joint effects of nutrient addition and enemy exclusion on exotic plant success.

Worldwide, ecosystems are increasingly dominated by exotic plant species, a shift hypothesized to result from numerous ecological factors. Two of these, increased resource availability and enemy release, may act in concert to increase exotic success in plant communities (Resource-Enemy Release Hypothesis, R-ERH). To test this, we manipulated the availability of soil nutrients and access of vertebrate herbivores, insect herbivores, and fungal pathogens to intact grassland communities containing both native and exotic species. Our results supported both conditions necessary for R-ERH. First, exotics were less damaged than natives, experiencing less foliar damage (insect herbivory and fungal disease) than native species, particularly in communities where soil nutrients were added. Second, fertilization increased foliar damage on native species, but not exotic species. As well as fulfilling both conditions for R-ERH, these results demonstrate the importance of considering the effects of resource availability when testing for enemy release. When both conditions are fulfilled, R-ERH predicts that increasing resource availability will increase exotic abundance only in the presence of enemies. Our results fully supported this prediction for vertebrate herbivores: fertilization increased exotic cover only in communities exposed to vertebrate herbivores. Additionally, the prediction was partially supported for insect herbivores and fungal pathogens, excluding these enemies reduced exotic cover as predicted, but inconsistent with R-ERH, this effect occurred only in unfertilized communities. These results highlight the need to consider the influence of multiple enemy guilds on community processes like exotic plant invasions. Moreover, this study experimentally demonstrates that resource availability and natural enemies can jointly influence exotic success in plant communities.

Reviews and syntheses: measuring ecosystem nitrogen status – a comparison of proxies

Abstract. There are many proxies used to measure nitrogen (N) availability in watersheds, but the degree to which they do (or do not) correlate within a watershed has not been systematically addressed. We surveyed the literature for intact forest or grassland watersheds globally, in which several metrics of nitrogen availability have been measured. Our metrics included the following: foliar δ15N, soil δ15N, net nitrification, net N mineralization, and the ratio of dissolved inorganic to organic nitrogen (DIN : DON) in soil solution and streams. We were particularly interested in whether terrestrial and stream based proxies for N availability were correlated where they were measured in the same place. Not surprisingly, the strongest correlation (Kendall's τ) was between net nitrification and N mineralization (τ = 0.71, p < 0.0001). Net nitrification and N mineralization were each correlated with foliar and soil δ15N (p < 0.05). Foliar and soil δ15N were more tightly correlated in tropical sites (τ = 0.68, p < 0.0001), than in temperate sites (τ = 0.23, p = 0.02). The only significant correlations between terrestrial- and water-based metrics were those of net nitrification (τ = 0.48, p = 0.01) and N mineralization (τ = 0.69, p = 0.0001) with stream DIN : DON. The relationship between stream DIN : DON with both net nitrification and N mineralization was significant only in temperate, but not tropical regions. To our surprise, we did not find a significant correlation between soil δ15N and stream DIN : DON, despite the fact that both have been used to infer spatially or temporally integrated N status. Given that both soil δ15N and stream DIN : DON are used to infer long-term N status, their lack of correlation in watersheds merits further investigation.

Soil carbon dioxide effluxes from different vegetation environments in semi-arid Eastern Cape, South Africa

Though important, CO2 effluxes from non-agricultural ecosystems are poorly characterised. Given that thicket vegetation is prevalent in Eastern Cape, South Africa, we monitored soil CO2 effluxes from an intact thicket, degraded thicket (DT) and grassland (G) over 10 months, as affected by temperature, moisture and penetration resistance (PR). High soil moisture (16%) reduced PR (4 kg cm−2), which raised winter effluxes (1.2 µmol m−2 s−1), while low moisture (2%) resulted in hard dry soil (14 kg cm−2) that suppressed spring effluxes (0.2 µmol m−2 s−1). There was good interaction between PR and moisture (r = −0.53), with seasonal effluxes increasing with increasing moisture (r = 0.9, p = 0.0001) and decreasing PR (r = −0.66, p = 0.02). Temperature effects were significant under unlimited moisture supply. Thus high summer temperature (40 °C) gave lower effluxes in DT and G (<1 µmol m−2 s−1) due to limited moisture (<10%), whereas high autumn temperature (48 °C) and good moisture (16%) accelerated CO2 emissions (averaging 2 µmol m−2 s−1) from all covers. Although semi-arid ecosystems are limited by low moisture and sandy soils, they contribute to CO2 emissions under high moisture and increasing temperature. Keywords: Eastern Cape, grassland, soil carbon dioxide effluxes, soil organic carbon, thicket

Landscape composition creates a threshold influencing Lesser Prairie-Chicken population resilience to extreme drought

Habitat loss and degradation compound the effects of climate change on wildlife, yet responses to climate and land cover change are often quantified independently. The interaction between climate and land cover change could be intensified in the Great Plains region where grasslands are being converted to row-crop agriculture concurrent with increased frequency of extreme drought events. We quantified the combined effects of land cover and climate change on a species of conservation concern in the Great Plains, the Lesser Prairie-Chicken (Tympanuchus pallidicinctus). We combined extreme drought events and land cover change with lek count surveys in a Bayesian hierarchical model to quantify changes in abundance of male Lesser Prairie-Chickens from 1978 to 2014 in Kansas, the core of their species range. Our estimates of abundance indicate a gradually decreasing population through 2010 corresponding to drought events and reduced grassland areas. Decreases in Lesser Prairie-Chicken abundance were greatest in areas with increasing row-crop to grassland land cover ratio during extreme drought events, and decreased grassland reduces the resilience of Lesser Prairie-Chicken populations to extreme drought events. A threshold exists for Lesser Prairie-Chickens in response to the gradient of cropland:grassland land cover. When moving across the gradient of grassland to cropland, abundance initially increased in response to more cropland on the landscape, but declined in response to more cropland after the threshold (δ=0.096, or 9.6% cropland). Preservation of intact grasslands and continued implementation of initiatives to revert cropland to grassland should increase Lesser Prairie-Chicken resilience to extreme drought events due to climate change.

Limiting processes for perennial plant reintroduction to restore dry grasslands

In restored grasslands of southern Europe, perennial plants remain highly underrepresented compared with the reference ecosystems. We tested various treatments to reintroduce common perennial plant species (Brachypodium retusum, Poaceae, and Thymus vulgaris, Lamiaceae), which are usually not or poorly reintroduced via soil and hay transfer. Treatments included microenvironmental manipulations (rock cover and plant interactions) and two grazing intensities. Target perennial species were transplanted in 2002 in the reference grassland ecosystem (intact grassland area used as a control) and in two abandoned fields. Survival was assessed in June 2003 and June 2004. Target species shoot and root biomass were measured in June 2004. Grazing greatly reduced the survival and biomass of both target species and its effects were reinforced by summer drought: plants that did not establish well enough during the autumn and spring did not survive summer. The restored rock cover had a mild positive effect, particularly on B. retusum. There were no negative or positive plant neighbor interactions in the steppe, while there was competition in both abandoned fields. Competition was particularly intense in the abandoned melon field, composed of a dense sward of annual grasses (Bromus sp.). In order to reintroduce perennial species to dry grasslands, the ideal combination of treatments is to exclude or reduce grazing during the first year to allow seedlings to establish and to recreate adequate microenvironmental conditions. Reducing competition from arable weeds may help but is not essential in such dry grasslands.

Assessing the status and conservation priorities of the short‐eared owl in North America

AbstractThe North American Breeding Bird Survey, Christmas Bird Count, and regional and national conservation assessments provide convincing evidence that the short‐eared owl (Asio flammeus) is experiencing a range‐wide, long‐term decline in abundance in North America. However, the species has received little conservation or research attention. The short‐eared owl is vulnerable to decline because it relies heavily on large, intact grasslands and a specialized diet of unpredictable small mammal prey. The species' nomadic movements compound these vulnerabilities by making a decline difficult to detect with current monitoring programs while obfuscating stewardship responsibilities for managers. The primary threat to the species is loss, fragmentation, and degradation of large tracts of native grasslands and wetlands. We propose the following conservation priorities to better understand and begin addressing the short‐eared owl's decline: 1) better define and protect important habitats; 2) improve population monitoring; 3) determine seasonal and annual movements; 4) re‐evaluate NatureServe's short‐eared owl national conservation classifications; 5) develop management plans and tools; and 6) classify raptors, including short‐eared owls, as migratory birds in Canada. We contend that the short‐eared owl's need for habitat conservation at large spatial scales, status as a predator, and high reproductive potential that affords the species capacity to recover, make it an effective and useful candidate as an umbrella species for grassland conservation. © 2014 The Wildlife Society.

Effect of aboveground litter on belowground plant interactions in a native Rough Fescue grassland

Chemical compounds from plants may exhibit stimulatory and/or inhibitory effects on surrounding organisms. However, research on belowground biochemical interactions among plants has focused more effort on elucidating negative effects. Moreover, the effect of shoot litter on belowground plant–plant interactions has remained relatively unexplored. In a field experiment with four target plant species (Artemisia frigida Willd., Solidago missouriensis Nutt.), Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths and Poa pratensis L.) interacting with intact grassland neighbours, we manipulated root competition using PVC tubes and shoot litter, and belowground chemical interaction by adding activated carbon (AC) to the soil. In A. frigida, shoot litter significantly interacted with root competition and root chemicals. Plants grown plus AC were larger than those minus AC when shoot litter was left intact suggesting inhibitory effects from neighbours and/or decomposing products. However, when shoot litter was removed, plants grown minus AC were larger suggesting stimulatory effects of root exudates. B. gracilis showed a similar trend but results were non-significant. Results demonstrate that the effects of neighbours can be inhibitory or facilitative depending on the presence or absence of shoot litter and mediation through AC.

Effect of chemical amendments to dairy soiled water and time between application and rainfall on phosphorus and sediment losses in runoff

Dairy soiled water (DSW) is a dilute, low nutrient effluent produced on Irish dairy farms through the regular washing down of milking parlours and holding areas. In Ireland, there is no closed period for the land application of DSW except where heavy rain is forecast within 48h. Chemical amendments have the potential to decrease phosphorus (P) and suspended sediment (SS) loss from DSW applied to land. This study examined the impact of three time intervals (12, 24 and 48h) between DSW application and rainfall and five treatments (control, unamended DSW, and DSW amended with lime, alum or ferric chloride (FeCl2)) on P and sediment losses from an intact grassland soil in runoff boxes. Rainfall was simulated at 10.5±1mmh−1. Phosphorus concentrations (1–1.6mgL−1) in runoff from DSW application, while not quantitative measures of P loss to surface waters in the field, indicated the importance of incidental P losses and that the current 48h restriction in Ireland is prudent. Unamended DSW application increased P loss by, on average, 71%, largely due to an increase in particulate phosphorus (PP) loss. All three amendments were effective in decreasing P and SS losses in runoff and, apart from the SS results for lime, were significantly different (p<0.05) to the control at at least one time point. Lime (a 64% reduction in total phosphorus (TP) in comparison with DSW only) was less effective than alum or FeCl2, likely due to the lower solubility of CaCO3 in water. Chemical amendment showed potential to decrease P losses from land application of DSW, but the efficacy of such amendments would need to be assessed in field trials and a cost–benefit analysis conducted to further examine whether they could be practically implemented on farms.

Density fractionation and 13C reveal changes in soil carbon following woody encroachment in a desert ecosystem

Woody encroachment has dramatically changed land cover patterns in arid and semiarid systems (drylands) worldwide over the past 150 years. This change is known to influence bulk soil carbon (C) pools, but the implications for dynamics and stability of these pools are not well understood. Working in a Chihuahuan Desert C4 grassland encroached by C3 creosote bush (Larrea tridentata), we used two density fractionation techniques (2 and 7 pool density fractionations) and isotopic analysis to quantify changes in C pools and dynamics among vegetation microsites typical of an encroachment scenario (remnant intact grassland, shrub subcanopies, and in shrub intercanopy spaces within a shrub-encroached area). The C concentration of bulk soils varied with microsite, with almost twice the C in shrub subcanopies as in intercanopy spaces or remnant grasslands. Estimated SOC accumulation rates from Larrea encroachment (4.79 g C m−2 year−1 under canopies and 1.75 g C m−2 year−1 when intercanopy losses were taken into account) were lower than reported for higher productivity Prosopis systems, but still represent a potentially large regional C sink. The composition of soil C varied among microsites, with the shrub subcanopy C composed of proportionally more light fraction C (<1.85 g cm−3) and C that was soluble in sodium polytungstate. Grassland soils contained very little carbonate C compared to shrub subcanopies or shrub intercanopy spaces. Stable isotope analyses indicate that inputs from C3 shrubs were incorporated into all density fractions, even in heavy fractions in which shrub inputs did not change overall C concentration. The seven density fractionation yielded unexpected δ13C patterns, where the two heaviest fractions were strongly depleted in 13C, indicating strong fractionation following organic matter inputs. These results suggest that the utility of isotope mixing models for determining input sources may be limited in systems with similar fractionation patterns. We propose a five pool model for dryland soil C that includes a relatively dynamic light fraction, aggregate and heavy fractions that are stable in size but that reflect dynamic inputs and outputs, a potentially large and seasonally dynamic pool of soluble C, and a large pool of carbonate C. Combined, these results suggest that dryland soil C pools may be more dynamic than previously recognized.

Evaluation of Amendments to Control Phosphorus Losses in Runoff from Pig Slurry Applications to Land

AbstractIf spread in excess of crop requirements, incidental phosphorus (P) losses from agriculture can lead to eutrophication of receiving waters. The use of amendments in targeted areas may help reduce the possibility of surface runoff of nutrients. The aim of this study was to identify amendments which may be effective in reducing incidental dissolved reactive phosphorus (DRP) losses in surface runoff from land applied pig slurry. For this purpose, the DRP losses under simulated conditions across the surface of intact grassland soil cores, loaded with unamended and amended slurry at a rate equivalent to 19 kg P ha−1, were determined over a 30 h period. The effectiveness of the amendments at reducing DRP in overlying water were (in decreasing order): alum (86%), flue gas desulfurization by‐product (FGD) (74%), poly‐aluminum (Al) chloride (PAC) (73%), ferric chloride (71%), fly ash (58%), and lime (54%). FGD was the most costly of all the treatments (€7.64/m3 for 74% removal). Ranked in terms of feasibility, which takes into account effectiveness, cost, and other potential impediments to use, they were: alum, ferric chloride, PAC, fly ash, lime, and FGD.

Vertebrate predators have minimal cascading effects on plant production or seed predation in an intact grassland ecosystem

The strength of trophic cascades in terrestrial habitats has been the subject of considerable interest and debate. We conducted an 8-year experiment to determine how exclusion of vertebrate predators, ungulates alone (to control for ungulate exclusion from predator exclusion plots) or none of these animals influenced how strongly a three-species assemblage of rodent consumers affected plant productivity. We also examined whether predator exclusion influenced the magnitude of post-dispersal seed predation by mice. Both ungulates and rodents had strong direct effects on graminoid biomass. However, rodent impacts on plant biomass did not differ across plots with or without predators and/or ungulates. Deer mice removed more seeds from seed depots on predator exclusion plots, suggesting trait-mediated indirect effects of predators, but this short-term behavioural response did not translate into longer-term impacts on seed survival. These results suggest that vertebrate predators do not fundamentally influence primary production or seed survival in our system.

A guilt-free roll in the grass: minimal short-term impacts from short-term camping in the Australian Alps

Back-country camping is a popular ecotourism activity in protected areas, including in Australia. Few studies have, however, used a manipulative experimental approach to quantify the impacts of camping. Here the impacts of four people camping for three nights on intact tall alpine herbfield and subalpine grasslands in the Australian Alps were quantified. Campsites were divided into three zones (tent, activity and control). Camping only caused decreases in vegetation height and cover in the tent and activity zones with only height effects still significant 6 weeks post-camping. This suggests that these alpine communities are relatively resistant to short-term camping activities by small groups. When the impacts of camping were compared with those from trampling experiments run concurrently, it is apparent that once these communities are damaged by recreation use, they are slower to recover. With consideration to this, dispersed and minimal impact practices should continue to be encouraged, as inappropriate behaviour (such as fire scars, cuts to trees, litter) and repeated campsite use may result in lasting damage.

Counterintuitive effects of large‐scale predator removal on a midlatitude rodent community

Historically, small mammals have been focal organisms for studying predator-prey dynamics, principally because of interest in explaining the drivers of the cyclical dynamics exhibited by northern vole, lemming, and hare populations. However, many small-mammal species occur at relatively low and fairly stable densities at temperate latitudes, and our understanding of how complex predator assemblages influence the abundance and dynamics of these species is surprisingly limited. In an intact grassland ecosystem in western Montana, USA, we examined the abundance and dynamics of Columbian ground squirrels (Spermophilus columbianus), deer mice (Peromyscus maniculatus), and montane voles (Microtus montanus) on 1-ha plots where we excluded mammalian and avian predators and ungulates, excluded ungulates alone, or allowed predators and ungulates full access. Our goal was to determine whether the relatively low population abundance and moderate population fluctuations of these rodents were due to population suppression by predators. Our predator-exclusion treatment was divided into two phases: a phase where we excluded all predators except weasels (Mustela spp.; 2002-2005), and a phase where all predators including weasels were excluded (2006-2009). Across the entire duration of the experiment, predator and/or ungulate exclusion had no effect on the abundance or overall dynamics of ground squirrels and deer mice. Ground squirrel survival (the only species abundant enough to accurately estimate survival) was also unaffected by our experimental treatments. Prior to weasel exclusion, predators also had no impacts on montane vole abundance or dynamics. However, after weasel exclusion, vole populations reached greater population peaks, and there was greater recruitment of young animals on predator-exclusion plots compared to plots open to predators during peak years. These results suggest that the impacts of predators cannot be generalized across all rodents in an assemblage. Furthermore, they suggest that specialist predators can play an important role in suppressing vole abundance even in lower-latitude vole populations that occur at relatively low densities.

Experimental manipulations of winter snow and summer rain influence ecosystem carbon cycling in a mixed‐grass prairie, Wyoming, USA

AbstractWater‐limited grasslands may be exceedingly vulnerable to changes in the timing or amount of precipitation which may result in shifts in the magnitudes and patterns of carbon cycling. Shifts in CO2 exchange may lead to alterations in carbon sequestration or net losses and could accentuate the rising CO2 concentrations in the atmosphere or ameliorate the increases. The objective of our project was to quantify how changes in winter, summer, and combined winter and summer precipitation may alter rates of ecosystem C cycling in the mixed‐grass prairie of the United States. Three replicated 50 m snow fences were installed to increase winter snow while summer precipitation was manipulated by either increasing (+50%) or decreasing (−50%) precipitation amounts. Measurements of net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (ER) and plant biomass were conducted throughout the snow‐free period. Deeper snow in winter increased summer ER by an average of 27%, GEP by 45%, NEE by 90% and plant biomass by 50% compared to ambient snow conditions. Average plant biomass increased 44% with the addition of summer rain and decreased by 18% where rainfall was excluded under ambient snow conditions. Average NEE was also lower where rain was excluded. Our findings indicate that seasonality changes of precipitation may be important in regulating the future extent of C sequestration and C cycling in one of the most extensive, intact grasslands of North America. Copyright © 2010 John Wiley &amp; Sons, Ltd.

Sod cutting and soil biota effects on seedling performance

Sod cutting (i.e. top soil removal) is a restoration management option for enhancing seedling establishment and for lowering the nutrient concentration in eutrophicated soils of nutrient-poor species-rich grasslands. Removal of the upper soil changes not only abiotic soil properties but may also affect the resident soil community. We investigated the effects of sod cutting on the establishment and performance of two endangered plant species ( Cirsium dissectum and Succisa pratensis) while simultaneously manipulating the interaction between seedlings and soil biota. In intact grassland and sod-cut areas at two localities, seedlings were grown in plastic tubes. Half of the tubes had a filter that excluded roots but allowed entry of fungal hyphae and soil microorganisms. The other tubes were closed (i.e. no contact with the surrounding soil). In a greenhouse experiment we studied the effect of soil solutions (with or without fungal tissue) from three grasslands and three sod-cut areas on seedling growth. Sod cutting had a positive net effect on seedling growth for S. pratensis. Access to (mycorrhizal) fungi and other soil biota resulted in a negative impact on seedling growth of both plant species, both in grassland and sod-cut areas. The greenhouse experiment confirmed that the soil biota in these meadows reduced seedling growth. Although sod cutting did not mitigate negative plant-soil feedback, it enhanced seedling growth, presumably by decreasing competition for light. Sod cutting is therefore very useful when seedling establishment needs to be stimulated.

MODEL ESTIMATION OF LAND-USE EFFECTS ON WATER LEVELS OF NORTHERN PRAIRIE WETLANDS

Wetlands of the Prairie Pothole Region exist in a matrix of grassland dominated by intensive pastoral and cultivation agriculture. Recent conservation management has emphasized the conversion of cultivated farmland and degraded pastures to intact grassland to improve upland nesting habitat. The consequences of changes in land-use cover that alter watershed processes have not been evaluated relative to their effect on the water budgets and vegetation dynamics of associated wetlands. We simulated the effect of upland agricultural practices on the water budget and vegetation of a semipermanent prairie wetland by modifying a previously published mathematical model (WETSIM). Watershed cover/land-use practices were categorized as unmanaged grassland (native grass, smooth brome), managed grassland (moderately heavily grazed, prescribed burned), cultivated crops (row crop, small grain), and alfalfa hayland. Model simulations showed that differing rates of evapotranspiration and runoff associated with different upland plant-cover categories in the surrounding catchment produced differences in wetland water budgets and linked ecological dynamics. Wetland water levels were highest and vegetation the most dynamic under the managed-grassland simulations, while water levels were the lowest and vegetation the least dynamic under the unmanaged-grassland simulations. The modeling results suggest that unmanaged grassland, often planted for waterfowl nesting, may produce the least favorable wetland conditions for birds, especially in drier regions of the Prairie Pothole Region. These results stand as hypotheses that urgently need to be verified with empirical data.

Biotic constraints on the invasion of diffuse knapweed (Centaurea diffusa) in North American grasslands

Knapweeds (Centaurea spp.) are among the most invasive of non-indigenous plant species that have colonized western North America over the last century. We conducted a 4-year experiment in a reconstructed grassland to test hypotheses related to the ability of grasslands to resist the invasion of diffuse knapweed (C. diffusa). We experimentally invaded C. diffusa and three native species into areas where we manipulated soil nitrogen (N) and phosphorus (P) availability and removed extant grasses to reduce competition. We evaluated the growth response of these species to these resources and competitive manipulations. Of the native species that were experimentally added, only one species, Ratibida pinnata (prairie coneflower), established in any numbers. Establishment values in intact vegetation were low for both species, but establishment by C. diffusa (0.02%) clearly outperformed that of R. pinnata (0.001%). Under reduced grass competition, establishment was enhanced, but the values for C. diffusa (0.68%) were not statistically different from those of R. pinnata (0.57%). Neither species performed better under higher soil nutrients in the presence of competing grasses. In plots with both species, biomass of the two planted species was positively correlated, but the biomass of both species was negatively correlated with non-added weedy species. Subsequent harvests of C. diffusa indicated that establishment was enhanced in treatments with higher soil nutrients but that the biomass of these plants could only be enhanced when plant competition was also reduced. These results indicate that C. diffusa can establish in intact grasslands at rates higher than natives, but opportunism rather than competitive ability best describes the invasiveness of C. diffusa. Thus, the mechanisms contributing to the establishment of this knapweed species are different from factors identified as contributing to the dominance of this invader.

The significance of small herbivores in structuring annual grassland

Abstract Question: Herbivores can play a fundamental role in regulating the composition and structure of terrestrial plant communities. Relatively inconspicuous but nevertheless ubiquitous gastropods and small mammals are usually considered to influence grassland communities through distinct modes. 1. Do terrestrial gastropods and small mammals, either alone or in combination, influence plant community composition of an intact annual grassland? 2. Do these herbivores influence the plant size structure of the dominant grass Avena? Location: Jasper Ridge Biological Preserve (37°24′ N, 122°13′ W, elevation 150 m) in northern California. Methods: Animal exclosures were used to examine the single and combined influences of these herbivores on annual grassland production, community composition, and plant size structure during the growing season of an intact annual grassland. Results: The removal and exclusion of the herbivores increased the prevalence of grasses relative to legumes and non-legume forbs; increas...

Phosphorus Leaching from Cow Manure Patches on Soil Columns

The loss of P in overland flow or leachate from manure patches can impair surface water quality. We studied leaching of P from 10-cm-high lysimeters filled with intact grassland soil or with acid-washed sand. A manure patch was created on two grassland and two sand-filled lysimeters, and an additional two grass lysimeters served as blanks. Lysimeters were leached in the laboratory during 234 d with a diluted salt solution, and column effluent was passed through a 0.45-microm filter, analyzed for pH, dissolved reactive P (DRP), and total dissolved P (TDP). At the end of the experiment lysimeter soil was sampled and analyzed for pH, available P, and oxalate-extractable P, Fe, and Al. The concentration of TDP in the effluent from the sand column increased to 25 mg L-1 during the first weeks and remained above 10 mg L-1 during the rest of the percolation. In effluent from grass + patch lysimeters TDP gradually increased to 4 mg L-1. Both in the manure and in the effluent of the sand lysimeter P was found mainly in the form of DRP, but in the effluent from the grass lysimeters was found mainly as dissolved unreactive P (DUP=TDP-DRP). Earthworm activity was responsible for decomposition of the manure patch on the grass lysimeters. Manure patches and their remains were found to be a long-term source of high concentrations of P in leachates. Spreading of patches after a grazing period could reduce their possible negative impacts on the environment.

Assessing grassland restoration success: relative roles of seed additions and native ungulate activities

Summary Grassland restorations often lack rare forb and grass species that are found in intact grasslands. The possible reasons for low diversity include seed limitation, microsite limitation and a combination of both. Native ungulates may create microsites for seedling establishment in tallgrass prairie restorations by grazing dominant species or through trampling activities, but this has never been tested in developing prairies. We experimentally tested for seed and microsite limitation in the largest tallgrass prairie restoration in the USA by adding rare forb and grass seeds in two trials inside and outside native ungulate exclosures. We measured seedling emergence because this stage is crucial in recruiting species into a community. We also measured light, water and standing crop biomass to test whether resource availability could help to explain seedling emergence rates. Ungulates increased light availability for each sampling time and also increased above‐ground net primary productivity (ANPP) during summer. Seedling emergence of rare prairie forbs and grasses was consistently greater when we added seeds. Seedling emergence was conditionally greater with a combination of seed additions and grazing, but grazing alone was unable to increase emergence. When ungulates increased seedling enhancement, the mechanism was partially associated with increased water and light availability. Exotic and cosmopolitan weed seedling emergence was not affected by grazing. Synthesis and applications. These results suggest that tallgrass prairie restorations are primarily seed limited and that grazing alone may not be able to increase seedling emergence of rare species without the addition of seeds. Therefore, adding seeds to grassland restorations may increase seedling emergence of rare species, and mimicking effects of grazing may increase emergence when seeds are added.