Respective influence of habitat conditions and management regimes on prealpine calcareous grasslands

The calcareous grasslands of the south-western French Alps have been poorly studied, although they provide suitable habitat for rare plant species and communities. The separate and combined effects on calcareous grassland communities of habitat conditions (lithology, soil moisture) and management regimes (grazing intensity, cutting regime) were studied using constrained ordination techniques (canonical correspondence analysis with variance partitioning). Among the explanatory variables considered, the most important factor determining floristic composition was lithology, which explained 11.9% of floristic variability, followed by grazing intensity (6.0%). Additive effects of management and lithology explained 23.9% of floristic variability. Species niche amplitude was measured by conditional variances of samples along main ordination axes, in order to define adequate conservation management for the rarest short-lived species with narrow niche breadth on both habitat and management gradients.     

Trampling Impacts on Thermotolerant Vegetation of Geothermal Areas in New Zealand

Geothermal features such as geysers, mud pools, sinter terraces, fumaroles, hot springs, and steaming ground are natural attractions often visited by tourists. Visitation rates for such areas in the Taupo Volcanic Zone of New Zealand are in the order of hundreds of thousands annually. These areas are also habitat for rare and specialized plant and microbial communities that live in the steam-heated soils of unusual chemical composition. We evaluated historical and current trampling impacts of tourists on the thermotolerant vegetation of the Waimangu and Waiotapu geothermal areas near Rotorua, and compared the results to experimental trampling at a third site (Taheke) not used by tourists. Historical tourism has removed vegetation and soil from around key features, and remaining subsoil is compacted into an impervious pavement on which vegetation recolonization is unlikely in the short term. Social tracks made by tourists were present at both tourist sites often leading them onto hotter soils than constructed tracks. Vegetation height and cover were lower on and adjacent to social tracks than further from them. Thermotolerant vegetation showed extremely low resistance to experimental trampling. This confirms and extends previous research that also shows that thallophytes and woody shrubs, life forms that dominate in thermotolerant vegetation, are vulnerable to trampling damage. Preservation of these vulnerable ecosystems must ensure that tourist traffic is confined to existing tracks or boardwalks, and active restoration of impacted sites may be warranted.     

Ground flora trampling studies: Five years after closure

Trampling as an ecological factor is a major concern of the management of park, forest, nature preserves, and wilderness areas as recreational resources. Of particular importance to the management concept of carrying capacity is the relative resistance of native plant communities, to trampling and the resilience or the capacity of impacted communities to recover. This information can be used by management to establish seasonal use limits to prevent irreversible degradation of these resources. The purpose of this study was to follow the recolonizaton of experimental trail surfaces barren of vegetative cover and hiked at three levels of use intensity. Results of this study indicate that soil compaction as measured by soil penetration resistance increased progressively with use level while the total number of species, species diversity, and dominant index scores declined. A major finding was that the greatest degree of change occurred at the first level of hiking, indicating that major floristic measures were most affected by the least amount of hiking. Recolonization of impacted areas that received 100 trampling passes as measured by plant cover, dominant indices, floristic dissimilarity, and species diversity was similar to areas receiving four and eight times more trampling, despite major differences in soil penetration resistance. These data suggest limited use delivered at one time can be as damaging as increasing levels of use delivered at over time.     

Characteristics of Urban Natural Areas Influencing Winter Bird Use in Southern Ontario,Canada

Characteristics of urban natural areas and surrounding landscapes were identified that best explain winter bird use for 28 urban natural areas in southern Ontario, Canada. The research confirms for winter birds the importance of area (size) and natural vegetation, rather than managed, horticultural parkland, within urban natural areas as well as percent urban land use and natural habitat in surrounding landscapes. Alien bird density and percent ground feeding species increased with percent surrounding urban land use. Higher percent forest cover was associated with higher percentages of forest, bark feeding, small (<20 g) and insectivorous species. Natural area size (ha) was related to higher species richness, lower evenness and higher percentages of insectivorous, forest interior, area-sensitive, upper canopy, bark feeding, and non-resident species. Higher number of habitat types within natural areas and percent natural habitat in surrounding landscapes were also associated with higher species richness. Common, resident bird species dominated small areas (<6.5 ha), while less common non-residents increased with area, indicative of a nested distribution. Areas at least 6.5 ha and more generally >20 ha start to support some area-sensitive species. Areas similar to rural forests had >25% insectivores, >25% forest interior species, >25% small species, and <5% alien species. Indicator species separated urban natural areas from rural habitats and ordination placed urban natural areas along a gradient between urban development and undisturbed, rural forests. More attention is needed on issues of winter bird conservation in urban landscapes.     

Responses of Three Heathland Shrubs to Single or Repeated Experimental Trampling

In the last few decades, trampling has become a cause of major disturbance to natural areas. Trampling can occur regularly either seasonally or throughout the year, or as a single impact depending on a recreational activity or an event. This study, carried out on a coastal heathland in north Brittany (France), was based on the hypothesis that the vegetation response can differ depending on whether the trampling is repeated or due to a single event. The aim was to compare the resistance and the short-term recovery capacity of selected species in coastal heathland communities subjected to a trampling experiment, applied once or five times (with equal total number of passes from 0 to 800). The temporal distribution of the trampling did not affect the response of Erica ciliaris, whereas for Erica cinerea and Ulex gallii, trampling was slightly less damaging when applied once rather than five times. These differences could be linked to the lower resistance and the higher recovery capacity of E. ciliaris, compared with the two other species. This study also emphasized the influence of site location by comparing observations with previous studies. Plant resistance calculated here appeared to be lower than that calculated for a less stressful site. The relationship between plant traits and response to trampling was also investigated. The effects of the different times between disturbance application and observations of the short-term response of woody vegetation to experimental trampling is also discussed.     

Effectiveness of Voluntary Habitat Stewardship in Conserving Grassland: Case of Operation Burrowing Owl in Saskatchewan

There have been no published performance evaluations of nongovernmental, voluntary habitat stewardship programs. The Operation Burrowing Owl (OBO) stewardship program, initiated in 1987, was evaluated for its effectiveness in conservation of grassland habitat during 1986–1993. The 108 OBO sites from 1987 to 88 and 98 randomly selected non-OBO sites that were grassland in 1986 in the Regina-Weyburn, Saskatchewan study area were classified by size and agricultural soil suitability. By 1993, 41 (38%) of the 108 OBO sites had been withdrawn from the program. The 1986 area of grassland was compared with grassland area calculated from digitized 1993 LANDSAT imagery. A correction for satellite inaccuracies was determined. Grassland retention in 1993 was significantly higher at OBO sites (66%) than at random sites (49%), demonstrating that the OBO voluntary program effectively conserved habitat. Also, grassland retention was significantly lower on sites with better agricultural soils, and for sites <12 ha in size. Site type (OBO or random), size and their interaction, followed by agricultural soil suitability, had the greatest effects on grassland retention. During an era of accelerated grassland loss, OBO strongly and positively (statistically significant) affected conservation of grassland sites most at risk: sites <12 ha in size and with good to excellent agricultural soils. This suggests that grassland conservation efforts focus on vulnerable sites (small size and/or good agricultural soils) to provide nesting habitat for burrowing owls. Our study demonstrates that a voluntary stewardship program can significantly increase conservation of habitat.     

Effects of Long-Term Trampling on the Above-Ground Forest Vegetation and Soil Seed Bank at the Base of Limestone Cliffs

Exposed limestone cliffs in central Europe harbor a highly divers flora with many rare and endangered species. During the past few decades, there has been increasing recreational use of these cliffs, which has caused local environmental disturbances. Successful restoration strategies hinge on identifying critical limitations. We examined the composition of aboveground forest vegetation and density and species composition of seeds in the soil seed bank at the base of four limestone cliffs in mixed deciduous forests that are intensively disturbed by human trampling and at four undisturbed cliffs in the Jura Mountains in northwestern Switzerland. We found that long-term human trampling reduced total aboveground vegetation cover at the base of cliffs and caused a significant shift in the plant-species composition. Compared with undisturbed cliffs, total seed density was lower in disturbed cliffs. Human trampling also altered the species composition of seeds in the soil seed bank. Seeds of unintentionally introduced, stress-tolerant, and ruderal species dominated the soil seed bank at the base of disturbed cliffs. Our findings indicate that a restoration of degraded cliff bases from the existing soil seed bank would result in a substantial change of the original unique plant composition. Active seed transfer, or seed flux from adjacent undisturbed forest areas, is essential for restoration success.     

Quantifying the effects of trampling and habitat edges on forest understory vegetation – A field experiment

We investigated the effects of human trampling on boreal forest understory vegetation on, and off paths from suburban forest edges towards the interiors and on the likelihood of trampling-aided dispersal into the forests for three years by carrying out a trampling experiment. We showed that the vegetation was highly sensitive to trampling. Even low levels of trampling considerably decreased covers of the most abundant species on the paths. Cover decreased between 10 and 30% on paths which had been trampled 35 times, and at least by 50% on those trampled 70–270 times. On-path vegetation cover decreased similarly at forest edges and in the interiors. However, some open habitat plant species that occurred outside the forest patches and at forest edges dispersed into the forests, possibly through the action of trampling. A higher cover percentage of an open habitat species at the forest edge line increased its probability to disperse into the forest interior. The vegetation community on, next to, and away from lightly trampled paths remained the same throughout the trampling experiment. For heavily trampled paths, the community changed drastically on the paths, but stayed relatively similar next to and away from the paths. As boreal vegetation is highly sensitive to the effects of trampling, overall ease of access throughout the forest floor should be restricted to avoid the excessive creation of spontaneous paths. To minimize the effects of trampling, recreational use could be guided to the maintained path network in heavily used areas.     

Restoring fire as an ecological process in shortgrass prairie ecosystems: initial effects of prescribed burning during the dormant and growing seasons

Prior to Anglo-European settlement, fire was a major ecological process influencing the structure, composition and productivity of shortgrass prairie ecosystems on the Great Plains. However during the past 125 years, the frequency and extent of grassland fire has dramatically declined as a result of the systematic heavy grazing by large herds of domestic cattle and sheep which reduced the available levels of fine fuel and organized fire suppression efforts that succeeded in altering the natural fire regime. The greatly diminished role of recurrent fire in these ecosystems is thought to be responsible for ecologically adverse shifts in the composition, structure and diversity of these grasslands, leading specifically to the rise of ruderal species and invasion by less fire-tolerant species. The purpose of this study was to evaluate the ecological effects of fire season and frequency on the shortgrass prairie and to determine the means by which prescribed fire can best be restored in this ecosystem to provide the greatest benefit for numerous resource values. Plant cover, diversity, biomass and nutrient status, litter cover and soil chemistry were measured prior to and following fire treatments on a buffalograss-blue grama shortgrass prairie in northeastern New Mexico. Dormant-season fire was followed by increases in grass cover, forb cover, species richness and concentrations of foliar P, K, Ca, Mg and Mn. Growing-season fire produced declines in the cover of buffalograss, graminoids and forbs and increases in litter cover and levels of foliar P, K, Ca and Mn. Although no changes in soil chemistry were observed, both fire treatments caused decreases in herbaceous production, with standing biomass resulting from growing-season fire approximately 600 kg/ha and dormant-season fire approximately 1200 kg/ha, compared with controls approximately 1800 kg/ha. The initial findings of this long-term experiment suggest that dormant-season burning may be the preferable method for restoring fire in shortgrass prairie ecosystems where fire has been excluded for a prolonged time period.     

A survey of trampling effects on vegetation and soil in eight tropical and subtropical sites

Impacts of recreation, especially of vehicles and walkers, were studied in eight tropical or subtropical public sites in Queensland. In each site, plant species number, vegetation cover, plant height, and species cover and frequency in untrampled, slightly trampled, moderately trampled, and heavily trampled areas were counted or measured. Soil penetration resistance and soil organic matter were also recorded. In two of these eight sites, plant cover, height, leaf length, leaf width, and leaf thickness of each species were measured. Some species of grass such asCynodon dactylon were present in areas subject to all degrees of trampling impact and some tussock species, particularlyEragrostis tenuifolia andSporobolus elongatus, were only present in trampled areas. Woody plants occurred only on untrampled areas. The number of species and all the vegetative measurements mentioned above were reduced as wear increased. Plant height was reduced dramatically by even light trampling. Tall plants appeared to be more sensitive to trampling than short plants. No clear relationship between soil organic matter content and trampling intensity was found.     

Scale-dependent inconsistencies in the effects of trampling on a forest understory community

The response of forest understory vegetation to trampling applied at different temporal and spatial scales was determined in a cliff-edge forest in Ontario, Canada. Three frequencies (0, 50, 500 passes per year) of short-term trampling (one year) were applied to plots previously undisturbed. Existing trails that had received three frequencies (approx. 100, 500, 25,000 passes per year) of long-term trampling (18 years) were also studied. Community composition, species richness, and individual species frequency were recorded in plots within 4 m and (or) 1 m of the patch centerline. The quantitative and qualitative form of plant response to increased trampling was compared for short-term and long-term treatments, both within 4 m and within 1 m of the path centerline, to judge the consistency of trampling effects at different temporal and spatial scales. As trampling frequency increased, community composition changed progressively, but consistently, in plots both within 4 m and 1 m of the path centerline. Species richness was less affected by trampling and only decreased within 1 m of the path centerline at the highest level of trampling (25,000 passes per season for 18 years). Effects of trampling on individual species frequency were much less consistent at different temporal and spatial scales of trampling. The scale-dependence results suggest that field workers and resource managers both should try explicitly to include and define multiple scale components when trying to ascertain the response of vegetation to human disturbance factors.     

Assessing the impacts of mountain biking and hiking on subalpine grassland in Australia using an experimental protocol

Mountain biking is an increasingly popular, but sometimes controversial, activity in protected areas. Limited research on its impacts, including studies comparing biking with hiking, contributes to the challenges for mangers in assessing its appropriateness. The impacts of mountain bike riding off trail were compared to those of hiking on subalpine grassland in Australia using a modification of a common trampling experimental methodology. Vegetation and soil parameters were measured immediately and two weeks after different intensities of mountain biking (none, 25, 75, 200 and 500 passes across slope, 200 pass up and down slope) and hiking (200 and 500 passes across slope). There were reductions in vegetation height, cover and species richness, as well as changes in species composition and increases in litter and soil compaction with riding. Riding up and down a moderate slope had a greater impact than riding across the slope. Hiking also affected vegetation height, cover and composition. Mountain biking caused more damage than hiking but only at high use (500 passes). Further research including other ecosystems, topography, styles of riding, and weather conditions are required, but under the conditions tested here, hiking and mountain biking appear to be similar in their environmental impacts.     

Seed bank composition of open and overgrown calcareous grassland soils--a case study from Southern Belgium

The success of calcareous grassland recreation following abandonment depends to a large extent on the composition of the soil seed bank. We studied the species richness and composition of the seed bank along a chronosequence from well-developed calcareous grassland to scrub and forest vegetation, which had developed on calcareous grassland over the last 225 years. The seed bank density was highest in the calcareous grassland soils (930 seeds/m(2)). However, the seed bank was mainly composed of rather common species of nutrient poor grassland, which were poorly represented in the surface vegetation. There were no significant differences in soil seed bank density and species composition between the calcareous grassland and scrub vegetation that was less than 15 years old, largely because several grassland species had persisted in the scrub vegetation and were therefore able to replenish the soil seed bank. In contrast seed density and species richness declined significantly after 40 years of grassland abandonment. Indeed, forest soils had the lowest seed densities (214 seeds/m(2)) with only a few grassland species represented. This reflects the lack of grassland species in the field layer of the forest, and, therefore, the lack of seed production and seed bank replenishment. It is clear that recreation of calcareous grassland on long abandoned sites cannot rely on germination of target species from the seed bank alone. Even in the calcareous grassland soil, seeds of target species are not abundant as the majority produces transient seeds, which decay rapidly if they do not germinate immediately. Successful grassland recreation on such sites therefore may require seeds of target species to be introduced artificially (e.g. as seed mixtures, green hay, etc.). Alternatively, reinstatement of traditional practices, such as grazing or mowing, will increase the natural dispersal potential of these species, allowing population reestablishment in the long term.     

Response of Three Plant Communities to Trampling in a Sand Dune System in Brittany (France)

Dunes that are protected because of their very rich and diverse plant communities are often exposed to excessive visitor pressure. The effects of trampling on the habitat must be known from a conservation viewpoint but also are important for management. To determine the response of plant assemblages to trampling by people, an experimental study was conducted on the state-owned dunes at Quiberon (Brittany, France). Indices of resistance and resilience were used to compare three typical plant communities belonging to the various landscape units: mobile dune, semifixed dune, and fixed dune. The strong contrasts between communities belonging to different successional stages reflect their ecological functioning. The mobile dune and semifixed dune with their low resistance contrasted with the fixed dune. Only the vegetation cover of the semifixed dune benefited from long-term trampling and had a very high resilience (134%). This response could be explained by a good balance of two opposite factors: soil compaction increasing soil stability and moisture content, and vegetation destruction. Because of their low resilience, trampling seems to be harmful for fixed dunes in the long term. The tourist pressure seems easier to integrate in to the mobile dunes and the semifixed dunes if periods of recovery are included in the management.     

Mediterranean Coastal Sand Dune Vegetation: Influence of Natural and Anthropogenic Factors

The aim of the present work was to assess the conservation status of coastal dune systems in Tuscany (Italy). Emphasis was given to the presence and abundance of plant communities identified as habitat in accordance with the Directive 92/43/EEC. Twenty transects perpendicular to the shoreline were randomly positioned on the whole coastal area (30 km in length) in order to sample the full spectrum of plant communities. Vegetation zonation and relationships with the most frequent disturbance factors in the study area—beach cleaning, coastline erosion, presence of paths and roads, bathing settlements and trampling—were investigated through principal coordinate analysis and canonical correspondence analysis. Natural factors, such as distance from the sea and total length, were also considered. Differences in the conservation status of the sites were found, ranging from the total disappearance of the foredune habitats to the presence of the complete psammophilous (sand-loving) plant communities. Erosion, trampling, and paths were found to be closely correlated with degradation and habitat loss. Furthermore, the overall plant species diversity of dunes was measured with NH_Dune, a modified version of the Shannon index; while the incidence of invasive taxa was calculated using N, a naturalness index. However, these diversity indices proved to be a weaker bioindicator of ecosystem integrity than habitat composition along transects. A possible strategy for the conservation and management of these coastal areas could be to protect the foredunes from erosion and limit trampling through the installation of footbridges or the use of appropriate fences.     

Estimation of Nitrous Oxide Emissions from US Grasslands

2 O) emissions from temperate grasslands are poorly quantified and may be an important part of the atmospheric N₂O budget. In this study N₂O emissions were simulated for 1052 grassland sites in the United States using the NGAS model of Parton and others (1996) coupled with an organic matter decomposition model. N₂O flux was calculated for each site using soil and land use data obtained from the National Resource Inventory (NRI) database and weather data obtained from NASA. The estimates were regionalized based upon temperature and moisture isotherms. Annual N₂O emissions for each region were based on the grassland area of each region and the mean estimated annual N₂O flux from NRI grassland sites in the region. The regional fluxes ranged from 0.18 to 1.02 kg N₂O N/ha/yr with the mean flux for all regions being 0.28 kg N₂O N/ha/yr. Even though fluxes from the western regions were relatively low, these regions made the largest contribution to total emissions due to their large grassland area. Total US grassland N₂O emissions were estimated to be about 67 Gg N₂O N/yr. Emissions from the Great Plains states, which contain the largest expanse of natural grassland in the United States, were estimated to average 0.24 kg N₂O N/ha/yr. Using the annual flux estimate for the temperate Great Plains, we estimate that temperate grasslands worldwide may potentially produce 0.27 Tg N₂O N/yr. Even though our estimate for global temperate grassland N₂O emissions is less than published estimates for other major temperate and tropical biomes, our results indicate that temperate grasslands are a significant part of both United States and global atmospheric N₂O budgets. This study demonstrates the utility of models for regional N₂O flux estimation although additional data from carefully designed field studies is needed to further validate model results.     

History Matters: Relating Land-Use Change to Butterfly Species Occurrence

Western European landscapes have drastically changed since the 1950s, with agricultural intensifications and the spread of urban settlements considered the most important drivers of this land-use/land-cover change. Losses of habitat for fauna and flora have been a direct consequence of this development. In the present study, we relate butterfly occurrence to land-use/land-cover changes over five decades between 1951 and 2000. The study area covers the entire Swiss territory. The 10 explanatory variables originate from agricultural statistics and censuses. Both state as well as rate was used as explanatory variables. Species distribution data were obtained from natural history collections. We selected eight butterfly species: four species occur on wetlands and four occur on dry grasslands. We used cluster analysis to track land-use/land-cover changes and to group communes based on similar trajectories of change. Generalized linear models were applied to identify factors that were significantly correlated with the persistence or disappearance of butterfly species. Results showed that decreasing agricultural areas and densities of farms with more than 10 ha of cultivated land are significantly related with wetland species decline, and increasing densities of livestock seem to have favored disappearance of dry grassland species. Moreover, we show that species declines are not only dependent on land-use/land-cover states but also on the rates of change; that is, the higher the transformation rate from small to large farms, the higher the loss of dry grassland species. We suggest that more attention should be paid to the rates of landscape change as feasible drivers of species change and derive some management suggestions.     

Reestablishing Naturally Functioning Dunes on Developed Coasts

Common beach management practices reduce the ecological values of coastal dunes. Mechanical beach cleaning eliminates incipient dunes, habitat for nesting birds, seed sources for pioneer dune colonizers and food for fauna, and artificially small, stabilized foredunes reduce the variability in microenvironments necessary for biodiversity. Recent initiatives for reducing coastal hazards, protecting nesting birds, and encouraging nature-based tourism provide incentive for the development of a restoration program for beaches and dunes that is compatible with human use. Suggested changes in management practice include restricting or rerouting pedestrian traffic, altering beach-cleaning procedures, using symbolic fences to allow for aeolian transport while preventing trampling of dunes, and eliminating or severely restricting exotic species. Landforms will be more natural in function and appearance but will be more dynamic, smaller and in a different position from those in natural areas. Research needs are specified for ecological, geomorphological, and attitudinal studies to support and inform restoration planning.     

The Impacts of Human Visitation on Mussel Bed Communities Along the California Coast: Are Regulatory Marine Reserves Effective in Protecting These Communities?

Rocky intertidal habitats frequently are used by humans for recreational, educational, and subsistence-harvesting purposes, with intertidal populations damaged by visitation activities such as extraction, trampling, and handling. California Marine Managed Areas, particularly regulatory marine reserves (MRs), were established to provide legal protection and enhancement of coastal resources and include prohibitions on harvesting intertidal populations. However, the effectiveness of MRs is unclear as enforcement of no-take laws is weak and no regulations protect intertidal species from other detrimental visitor impacts such as trampling. The goal of this study was two-fold: (1) to determine impacts from human visitation on California mussel populations (Mytilus californianus) and mussel bed community diversity; and (2) to investigate the effectiveness of regulatory MRs in reducing visitor impacts on these populations. Surveys of mussel populations and bed-associated diversity were compared: (1) at sites subjected to either high or low levels of human use, and (2) at sites either unprotected or with regulatory protection banning collecting. At sites subjected to higher levels of human visitation, mussel populations were significantly lower than low-use sites. Comparisons of mussel populations inside and outside of regulatory MRs revealed no consistent pattern suggesting that California no-take regulatory reserves may have limited effectiveness in protecting mussel communities. In areas where many people visit intertidal habitats for purposes other than collecting, many organisms will be affected by trampling, turning of rocks, and handling. In these cases, effective protection of rocky intertidal communities requires an approach that goes beyond the singular focus on collecting to reduce the full suite of impacts.