Barriers to shrub reestablishment following fire in the seasonal submontane zone of Hawai'i

Introduced grass species have invaded extensive areas of Hawaii Volcanoes National Park and increased the size and frequency of fire. Following fire, grass cover is enhanced while native shrub cover is reduced; the reduction in most shrubs persists for at least 20 years even in the absence of fire. Shrub seedlings were planted in burned and unburned plots with and without grass cover. Biomass of 14 month old shrub seedlings was generally highest in recently burned/grass removed plots, intermediate in old burn/grass removed plots, and lowest in unburned/grass removed plots. In contrast, shrub biomass in plots with grass cover was low and did not differ significantly among burn treatments. Light competition is likely to be responsible for differences in shrub growth rates; grass cover reduced light to 1–10% of background levels. In addition, pool sizes of available soil N were highest in recently burned, intermediate in old burn, and lowest in unburned areas.     

Mortality of native grasses after a summer fire in natural temperate grassland suggests ecosystem instability

The exclusion of regular fire and the introduction of livestock grazing have altered native grassland composition on Victoria's volcanic plains, commonly resulting in spear‐grass and wallaby‐grass pastures replacing Kangaroo Grass grasslands. The effect of reintroducing fire to these pastures is currently unknown, although it may be an important part of restoring this ecosystem. We measured the changes in basal area of the dominant grasses in a mixed Spear‐grass/Wallaby‐grass pastures after a summer wildfire, which we assume burnt a relatively homogenous grass sward. We found a 90–95% reduction in the basal area of live spear‐grass tussocks in burnt plots compared with unburned controls, due to the mortality of tussocks. This suggests that caution and structured experimentation should be applied when using fire to manage spear‐grass‐dominated grasslands.     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

Patterns of herbaceous plant diversity in southeastern Louisiana pine savannas

Question: How diverse are Louisiana pine savanna plant communities and how is diversity affected by time since burn and removal of a competitively dominant species? Location: Lake Ramsay, southeastern Louisiana, USA. Methods: Species‐area curves were constructed from nine nested quadrats in open savanna differing in time since burn (6, 18 and 30 months). Species frequency was determined for 100 1 ‐m² quadrats. The dominant gras s, Andropogon virginicus, was removed with herbicide from moist and dry sites to test for possible effects of competition. Results: Slopes of log‐log species‐area relationships were steep (0.195 to 0.379). Time since burn did not affect the richness of herbaceous plants, only woody species. More than half of all species recorded (43/79, 54%) were infrequent (in < 10% of quadrats). After two years, there were no differences in species richness and composition for plots with and without A. virginicus. Conclusions: The high species diversity is typical of other savannas across the coastal plain. The large number of infrequent species indicates that the core‐satellite pattern of species occurrence found in temperate grasslands does not apply to southern pine savannas. The absence of effects due to removal of a dominant may be due to insufficient observation time, or low competition. Most species have traits, such as diminutive life forms, that suggest they are weak competitors for light in the presence of robust matrix grasses and in the absence of fire. Many species in Pinuspalustris savannas are likely either fugitive or peripheral species.     

Soil seed bank dynamics of fire-prone wooded grassland, woodland and dry forest ecosystems in Ethiopia

In order to reveal the role of soil seed banks in vegetation recovery after fire in savanna, the spatial distribution and temporal changes in the soil seed banks of regularly burning savanna in Gambella, western Ethiopia, was studied. The seedling emergence technique was employed to determine the species composition and density of the soil seed bank of six sites ranging in fire severity from wooded grassland with frequent fires over woodland with intermediate fire frequency to forest with absence of fires. Species composition and density of seeds in the soil were compared between seasons, depths and sites with different types of standing vegetation. Fourteen plant species were recorded in the soil seed bank from the grassland and woodland sites and 6 from the dry forests; 60 % of the taxa in the soil seed bank were annuals and 40 % were perennials. The soil seed banks were largely dominated by graminoids and 48–97 % of the soil seed bank in the grasslands and woodlands was of a single grass species, Hyparrhenia confinis , which was absent from the dry forests. The soil seed pools ranged from less than 100 to 4700 seeds per m² depending upon the season. The soil seed bank of graminoids was nearly empty after the onset of the rainy season whereas seeds of broadleaved herbs and woody species able to germinate were still found after this time. Floristic composition, representation of life forms and density of seeds in soil did not correspond closely with that of the standing vegetation, but within graminoids there was a strong similarity between the soil seed bank and the standing vegetation. The current fire regime of Ethiopian savanna woodlands appears to maintain the dominance of graminoids over broadleaved herbs and woody plants both as seeds in the soil and in the standing vegetation.     

The effect of prescribed burning on plant rarity in a temperate forest

Rare species can play important functional roles, but human‐induced changes to disturbance regimes, such as fire, can inadvertently affect these species. We examined the influence of prescribed burns on the recruitment and diversity of plant species within a temperate forest in southeastern Australia, with a focus on species that were rare prior to burning. Floristic composition was compared among plots in landscapes before and after treatment with prescribed burns differing in the extent of area burnt and season of burn (before–after, control‐impact design). Floristic surveys were conducted before burns, at the end of a decade of drought, and 3 years postburn. We quantified the effect of prescribed burns on species grouped by their frequency within the landscape before burning (common, less common, and rare) and their life‐form attributes (woody perennials, perennial herbs or geophytes, and annual herbs). Burn treatment influenced the response of rare species. In spring‐burn plots, the recruitment of rare annual herbs was promoted, differentiating this treatment from both autumn‐burn and unburnt plots. In autumn‐burn plots, richness of rare species increased across all life‐form groups, although composition remained statistically similar to control plots. Richness of rare woody perennials increased in control plots. For all other life‐form and frequency groups, the floristic composition of landscapes changed between survey years, but there was no effect of burn treatment, suggesting a likely effect of rainfall on species recruitment. A prescribed burn can increase the occurrence of rare species in a landscape, but burn characteristics can affect the promotion of different life‐form groups and thus affect functional diversity. Drought‐breaking rain likely had an overarching effect on floristic composition during our study, highlighting that weather can play a greater role in influencing recruitment and diversity in plant communities than a prescribed burn.     

Fire effects on mycorrhizal symbiosis and root system architecture in southern African savanna grasses

Mycorrhizal symbiosis is a key factor influencing aspects of grassland and savanna structure and functioning including plant growth, competition, population and community dynamics, and responses to fire and herbivory. This study assessed the effects of fire on mycorrhizal symbiosis and root system architecture (RSA) in South African savanna grasses. Eighteen grass species were sampled across contrasting fire frequency treatments in the Kruger National Park experimental burn plots. All eighteen species studied were highly colonized by arbuscular mycorrhizal fungi (AMF). Both mycorrhizal symbiosis and RSA were strongly affected by fire, with an increase in AMF colonization and a decrease in root branching and fine root development with decreasing fire frequency. Greater water limitation in frequently burned savanna may result in greater fine root development, thus reducing plant dependency on AMF for acquisition of soil resources. Reduced mycorrhizal colonization in frequently burned savanna may also be driven by higher phosphorus : nitrogen ratios, or indirect effects related to higher grazing intensities in frequently burned sites.     

Assessment of woody species encroachment in the grasslands of Nechisar National Park,Ethiopia

Ecological survey was executed to assess woody species encroachment into the grassland plain of Nechisar National Park (NNP). Forty‐one woody species were recorded. Dichrostachys cinerea Wight & Arn., Acacia mellifera (Vahl) Benth., Acacia nilotica (L) Willd., Acacia senegal (L.) Willd., Acacia seyal Del. and Acacia tortilis (Forssk.) Hayne were among the major encroaching woody species. The majority of the woody species were found to be highly aggregated in their pattern of distribution, while only few species showed some degree of randomness. The mean woody species density was ca. 1995 woody plants ha^?1. Mean cover of woody, grass, unpalatable forbs and total herbaceous species were 31%, 58%, 68% and 121%, respectively. The woody species density and cover, unpalatable forbs and bare land cover were significantly higher in the highly grazed and fire‐suppressed part of the grassland plain. Pearson correlation coefficient matrix indicated that woody species cover and density were negatively correlated with total herbaceous and grass cover. The high woody, unpalatable forbs and bare land cover indicated the progressively increasing perennial grass species diversity deterioration in the grass plain of the Park. Decline in the grassland condition, unless reversed, will jeopardize the biological diversity as well as the aesthetic value of the NNP.     

Effects of experimental season of prescribed fire and nutrient addition on structure and function of previously grazed grassland

Aims Understanding the drivers of grassland structure and function following livestock removal will inform grassland restoration and management. Here, we investigated the effects of fire and nutrient addition on structure and function in a subtropical semi-native grassland recently released from grazing in south-central Florida. We examined responses of soil nutrients, plant tissue nutrients, biomass of live, standing dead and litter, and plant species composition to experimental annual prescribed fire applied during different seasons (wet season vs. dry season), and nutrient additions (N, P and N + P) over 9 years.Methods Experimental plots were set up in a randomized block split-plot design, with season of prescribed fire as the main treatment and nutrient addition as the subplot treatment. Species cover data were collected annually from 2002 to 2011 and plant tissue and plant biomass data were collected in 2002–2006 and 2011. Soil nutrients were analyzed in 2004, 2006 and 2011.Important findings Soil total phosphorus (P) levels increased substantially with P addition but were not influenced by prescribed fire. Addition of P and N led to increased P and N concentrations in live plant tissues, but prescribed fire reduced N in live tissue. Levels of tissue N were higher in all plots at the beginning of the experiment, an effect that was likely due to grazing activity prior to removal of livestock. Plant tissue N steadily declined over time in all plots, with annually burned plots declining faster than unburned plots. Prescribed fire was an important driver of standing dead and litter biomass and was important for maintaining grass biomass and percent cover. Nutrient addition was also important: the addition of both N and P was associated with greater live biomass and woody forbs. Removal of grazing, lack of prescribed fire, and addition of N + P led to a reduction of grass biomass and a large increase in biomass of a woody forb. Annual prescribed fire promoted N loss from the system by reducing standing dead and litter, but maintained desirable biomass of grasses.     

Long-term patterns of shrub expansion in a C4-dominated grassland: fire frequency and the dynamics of shrub cover and abundance

Worldwide, grassland ecosystems have experienced a major shift in growth-form dominance as woody plant species have expanded and replaced native grasses. In the C(4)-dominated grasslands of central North America, a reduction in fire frequency is the most cited cause of this shift in growth forms as fire both enhances grass productivity and constrains the establishment and expansion of native woody vegetation. Using an 18-yr plant species composition data set, we quantified patterns of change in shrub cover, frequency, and species richness associated with three distinct fire regimes. During the study period (1983-2000), shrub cover increased most dramatically in sites in which the frequency of fire was once every 4 yr (intermediate frequency; 28.6%) followed by sites in which fire occurred only once during the 18-yr period (low frequency; 23.7%). Annual fire effectively prevented the recruitment of new woody species, but even with this high fire frequency, shrub cover increased slightly (3.7%). Comparatively, shrub species richness increased by three and six, respectively, in the intermediate- and low-frequency fire sites. These data indicate that within this grassland, periods without fire are necessary for recruitment of both new individuals and additional shrub species; however, once established, shrub cover will increase regardless of fire frequency and even annual fire will not reduce shrub abundance.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.     

Invertebrate community response to fire and rodent activity in the Mojave and Great Basin Deserts

Recent increases in the frequency and size of desert wildfires bring into question the impacts of fire on desert invertebrate communities. Furthermore, consumer communities can strongly impact invertebrates through predation and top‐down effects on plant community assembly. We experimentally applied burn and rodent exclusion treatments in a full factorial design at sites in both the Mojave and Great Basin deserts to examine the impact that fire and rodent consumers have on invertebrate communities. Pitfall traps were used to survey invertebrates from April through September 2016 to determine changes in abundance, richness, and diversity of invertebrate communities in response to fire and rodent treatments. Generally speaking, rodent exclusion had very little effect on invertebrate abundance or ant abundance, richness or diversity. The one exception was ant abundance, which was higher in rodent access plots than in rodent exclusion plots in June 2016, but only at the Great Basin site. Fire had little effect on the abundances of invertebrate groups at either desert site, with the exception of a negative effect on flying‐forager abundance at our Great Basin site. However, fire reduced ant species richness and Shannon's diversity at both desert sites. Fire did appear to indirectly affect ant community composition by altering plant community composition. Structural equation models suggest that fire increased invasive plant cover, which negatively impacted ant species richness and Shannon's diversity, a pattern that was consistent at both desert sites. These results suggest that invertebrate communities demonstrate some resilience to fire and invasions but increasing fire and spread of invasive due to invasive grass fire cycles may put increasing pressure on the stability of invertebrate communities.     

Fire,aridity and seed banks. What does seed bank composition reveal about community processes in fire‐prone desert?

Questions: The relationship between fire, aridity and seed banks is poorly understood in plant community ecology. We tested whether there was a close correspondence between the seed bank and standing vegetation composition with time‐since‐fire in a desert. We also examined whether longer‐lived species showed seed limitation relative to more ephemeral species, as this could influence grass‐woody ratios in a major biome. Location: Dune hummock grasslands/shrublands of central Australia. Methods: The effects of time‐since‐fire on floristic and functional group composition were examined by comparing plots unburned since 1984 against plots that had been burned in 2002. Three methods were used to quantify seed abundances: a germination trial using heat and smoke application, a flotation method, and a sieving method. Results: Seed bank densities were very low (<3000 m^?2). Species similarity between the seed bank and standing vegetation was high at sites recently burned (0.86) and low in sites long‐since burned (0.52). The relative abundance of ephemeral species in the seed bank peaked in recently burned plots, but the relative abundance of seeds of woody species did not match the pattern of abundance in the standing vegetation. Remarkably, the dominant perennial grasses and woody species were either absent from the seed bank or present at extremely low abundances. Discussion: Differences in the relative abundance of ephemeral species between standing vegetation and seed bank relate to the post‐fire succession process. The small soil pool of seed from woody species may be explained by allocation to belowground carbohydrate storage over seed production. Field observations suggest, however, that production of strongly dormant seed can be prolific and that high levels of seed predation make this system strongly seed‐limited. The discovery of this seed bank syndrome indicates that shifts in grass‐woody ratios can be driven by the juxtaposition of unpredictable seed rain and fire events in these desert dunes. However, estimates of grass‐woody ratios due to changing fire regimes will be difficult to predict.     

Fire-mediated dieback and compositional cascade in an Amazonian forest

The only fully coupled land-atmosphere global climate model predicts a widespread dieback of Amazonian forest cover through reduced precipitation. Although these predictions are controversial, the structural and compositional resilience of Amazonian forests may also have been overestimated, as current vegetation models fail to consider the potential role of fire in the degradation of forest ecosystems. We examine forest structure and composition in the Arapiuns River basin in the central Brazilian Amazon, evaluating post-fire forest recovery and the consequences of recurrent fires for the patterns of dominance of tree species. We surveyed tree plots in unburned and once-burned forests examined 1, 3 and 9 years after an unprecedented fire event, in twice-burned forests examined 3 and 9 years after fire and in thrice-burned forests examined 5 years after the most recent fire event. The number of trees recorded in unburned primary forest control plots was stable over time. However, in both once- and twice-burned forest plots, there was a marked recruitment into the 10-20cm diameter at breast height tree size classes between 3 and 9 years post-fire. Considering tree assemblage composition 9 years after the first fire contact, we observed (i) a clear pattern of community turnover among small trees and the most abundant shrubs and saplings, and (ii) that species that were common in any of the four burn treatments (unburned, once-, twice- and thrice-burned) were often rare or entirely absent in other burn treatments. We conclude that episodic wildfires can lead to drastic changes in forest structure and composition, with cascading shifts in forest composition following each additional fire event. Finally, we use these results to evaluate the validity of the savannization paradigm.     

Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

Effects of kangaroo rat exclusion on vegetation structure and plant species diversity in the Chihuahuan Desert

Long-term (1977–90) experimental exclusion of three species of kangaroo rats from study plots in the Chihuahuan Desert resulted in significant increases in abundance of a tall annual grass (Aristida adscensionis) and a perennial bunch grass (Eragrostis lehmanniana). This change in the vegetative cover affected use of these plots by several other rodent species and by foraging birds. The mechanism producing this change probably involves a combination of decreased soil disturbance and reduced predation on large-sized seeds when kangaroo rats are absent. Species diversity of summer annual dicots was greater on plots where kangaroo rats were present, as predicted by keystone predator models. However, it is not clear whether this was caused directly by activities of the kangaroo rats or indirectly as a consequence of the increase in grass cover. No experimental effect on species diversity of winter annual dicots was detected. Our study site was located in a natural transition between desert scrub and grassland, where abiotic conditions and the effects of organisms may be particularly influential in determining the structure and composition of vegetation. Under these conditions kangaroo rats have a dramatic effect on plant cover and species composition.     

Effects of invasion of fire‐free arid shrublands by a fire‐promoting invasive alien grass (Pennisetum setaceum) in South Africa

Arid shrublands in the Karoo (South Africa) seldom accumulate sufficient combustible fuel to support fire. However, as a result of invasion by an alien perennial grass (Pennisetum setaceum), they could become flammable. This paper reports on an experiment to assess the effects of fire following invasion by P. setaceum. We established 10 plots (5 × 10 m) separated by 2.5 m, and added grass fuel to five plots (5 and 10 tons ha^?1 to alternate halves of the plot) leaving the remaining five plots as interspersed controls. Plots with fuel added were burnt, and fire behaviour was measured during the burns. Rates of fire spread were generally low (0.01–0.07 m s^?1) and did not differ significantly between burn treatments. Mean fireline intensities were higher in the high compared with the low fuel treatments (894 and 427 kW m^?1, respectively). We recorded plant species and their cover before and after burning on each of the plots. After 15 months of follow‐up monitoring in the burn plots, only two species, the dwarf shrub (Tripteris sinuata) and the perennial herb (Gazania krebsiana) resprouted. Most individuals of other species were killed and did not reseed during the 15‐month study. The mass of added fuel load (high or low) did not influence vegetation recovery rates after fire. Should future invasions by P. setaceum lead to similar fuel loads in these shrublands, inevitable fires could change the vegetation and may favour spread of the flammable grass. Our results have important implications for predicting the effects of invasive alien plants (especially grasses) on fire‐free ecosystems elsewhere. The predicted impacts of fire may alter species composition, ultimately affecting core natural resources that support the Karoo economy.     

Species richness and vegetation structure in a limestone grassland after 15 years management with six biomass removal regimes

Species richness, species composition and standing crop were studied in a limestone grassland in northern Switzerland after 13 and 15 years of experimental management with 6 management regimes differing in form, time and frequency of biomass removal. The management regimes with a cut at least every 2nd year had a higher species number per 40 m² and 1 m² than those cut less frequently, abandoned or annually burned. Annual burning and a cut every 5th year lead to a similar decrease in species richness as abandonment. Differences in species richness between different regimes increased with increasing spatial resolution of observation. Many species occurred in a plot during only one of the two sample years. This turnover of species was the highest in plots with the lowest species numbers, and increased with increasing spatial resolution. Standing crop was highest in plots burned annually (290 g m^?2), and lowest in plots cut annually in July (1989 g m^?2). Standing crop and litter increased with decreasing frequency of cutting. *** DIRECT SUPPORT *** A02DO006 00005     

Fire as a Recurrent Event in Tropical Forests of the Eastern Amazon: Effects on Forest Structure,Biomass, and Species Composition1

The effects of fire on forest structure and composition were studied in a severely fire-impacted landscape in the eastern Amazon. Extensive sampling of area forests was used to compare structure and compositional differences between burned and unburned forest stands. Burned forests were extremely heterogeneous, with substantial variation in forest structure and fire damage recorded over distances of <50 m. Unburned forest patches occurred within burned areas, but accounted for only six percent of the sample area. Canopy cover, living biomass, and living adult stem densities decreased with increasing fire inrensiry / frequency, and were as low as 10–30 percent of unburned forest values. Even light burns removed >70 percent of the sapling and vine populations. Pioneer abundance increased dramatically with burn intensity, with pioneers dominating the understory in severely damaged areas. Species richness was inversely related to burn severity, but no clear pattern of species selection was observed. Fire appears to be a cyclical event in the study region: <30 percent of the burned forest sample had been subjected to only one burn. Based on estimated solar radiation intensities, burning substantially increases fire susceptibility of forests. At least 50 percent of the total area of all burned forests is predicted to become flammable within 16 rainless days, as opposed to only 4 percent of the unburned forest. In heavily burned forest subjected to recurrent fires, 95 percent of the area is predicted to become flammable in <9 rain-free days. As a recurrent disturbance phenomenon, fire shows unparalleled potential to impoverish and alter the forests of the eastern Amazon.     

Long-term after effects of fertilization on above-ground phytomass and species diversity in calcareous grassland

Abstract. Long-term after effects on species number and productivity in calcareous grassland were analyzed after cessation of fertilization. Three series of permanent plots were monitored yearly from 1971 to 1993. These series differed in duration of fertilization and in fertilizer composition, notably the amount of nitrogen and phosphorus. Yearly above-ground production decreased in all series after fertilization had been stopped, however at different speeds. The grass/forb ratio also decreased, while species number and Shannon index of diversity (H') increased. In the series where fertilizer treatment was stopped in 1967 (1968-series), productivity and grass/forb ratio decreased equally as in the plots where fertilization with a high nitrogen content was stopped in 1979 (80/Npk-series). However, species number in the plots of the 80/Npk-series increased faster than in the 1968-series. This was probably the result of a higher number of species present in the seed rain from the surrounding vegetation in the period after 1980 than from after 1968. In the plots receiving a high amount of phosphorus (80/nPk series), the productivity decreased more slowly than in the high nitrogen plots (80/Npk-series). At approximately the same total production level, the grass/forb ratio remained higher for another five-year period in the 80/nPk-series. Species number and the Shannon index of diversity increased more slowly in the 80/nPk-series than in the 80/Npk-series. Over a given range of productivity, changes in species number and Shannon index correlated better with the grass/forb ratio than with total above-ground phytomass. Therefore, restoration of species-rich grassland should not only be focused on lowering the yearly production, but also on reducing the grass component of the vegetation.