Variation in nitrogen deposition and available soil nitrogen in a forest–grassland ecotone in Canada

Regional variation in nitrogen (N) deposition increases plant productivity and decreases species diversity, but landscape- or local -scale influences on N deposition are less well-known. Using ion-exchange resin, we measured variation of N deposition and soil N availability within Elk Island National Park in the ecotone between grassland and boreal forest in western Canada. The park receives regionally high amounts of atmospheric N deposition (22 kg ha⁻¹ yr⁻¹). N deposition was on average higher ton clay-rich luvisols than on brunisols, and areas burned 1–15 years previously received more atmospheric N than unburned sites. We suggest that the effects of previous fires and soil type on deposition rate act through differences in canopy structure. The magnitude of these effects varied with the presence of ungulate grazers (bison, moose, elk) and vegetation type (forest, shrubland, grassland). Available soil N (ammonium and nitrate) was higher in burned than unburned sites in the absence of grazing, suggesting an effect of deposition. On grazed sites, differences between fire treatments were small, presumably because the removal of biomass by grazers reduced the effect of fire. Aspen invades native grassland in this region, and our results suggest that fire without grazing might reinforce the expansion of forest into grassland facilitated by N deposition.     

Scales of movement by elk (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in response to heterogeneity in forage resources and predation risk

Animals may respond to spatial and temporal heterogeneity by altering their movement patterns. The time an animal spends in an area of a given size is termed ȁ8first-passage timeȁ9 and can be used to identify the scales at which different movement processes occur. Using first-passage time and 2-h observations, we identified nested spatial scales representing three movement behaviours for elk (Cervus elaphus) – inactive/resting (moves < 50 m), active/foraging (x̄ = 276.7 m, SD = 56.6), and active/relocating (x̄ = 1628.3 m, SD = 436.6). Our ability to identify inactive behaviour was limited by GPS accuracy. The scale separating relocating and foraging behaviour ranged 550–1650 m across individuals and varied quadratically with the mean patch size of cutover forest in an animal’s home range. We classified path segments into the 3 movement behaviours and related behaviours to local environmental conditions. Elk were likely to be inactive in areas having a low predicted use by wolves (Canis lupus), farther than 50 m from anthropogenic linear clearings, and where microclimatic conditions were cool (high shrub cover and north to east-facing slopes). In contrast, elk were most likely to forage in areas having intermediate levels of herbaceous biomass and low movement costs. Elk were most likely to be relocating when in areas of high wolf use, when close to linear clearings, and in energetically costly situations such as moving upslope. We discuss how elk use of potential foraging habitats may be restricted in this landscape by risks imposed by predators, humans, or both.     

The spatial extent and relative influence of landscape-level factors on wintering bird populations

The influences of the landscape matrix (complex of habitats surrounding a study plot) and within-patch vegetation were studied in bird communities wintering in the piedmont of Georgia, USA. Variation at the landscape and within-patch levels was controlled to reduce the likelihood of confounding and spurious relationships. The landscape matrix within 500 m of each study plot was quantified from aerial photographs. Statistical models using landscape matrix and within-patch vegetation variables explained 73–84% of variation in bird abundance and diversity among sites with landscape matrix variables accounting for 30–90% of the variation. Variation in bird species richness and diversity was explained solely by landscape variables. Models for individual species such as Carolina Wrens (Thyrothorus ludovicianus) and Rufous-sided Towhees (Pipilo erythrophthalmus) had r² > 0.80, with the landscape matrix variables accounting for the majority of this variation. However, other species like Northern Cardinals (Cardinalis cardinalis) and White-throated Sparrows (Zonotrichia albicollis) were most strongly influenced by within-plot vegetation. The landscape influence extended beyond habitats immediately adjacent to the study plots as indicated by significant variables describing variation in more distant habitat patches. These analyses illustrate a technique for comparing the strength of within-patch versus landscape influences and measuring the spatial extent of the landscape influence in fine-grained landscapes.Report No. 3955, Environmental Sciences Division, Oak Ridge National Laboratory.This research received funding from the Ecological Research Division, Office of Health and Environmental Research, U.S. Department of Energy, under Contract No. DE-AC05-84OR21400 with Martin Marietta Energy Systems, Inc.     

The effect of spatial scale on Konza landscape classification using textural analysis

Spatial scale is inherent in the definition of landscape heterogeneity and diversity. For example, a landscape may appear heterogeneous at one scale but quite homogeneous at another scale. In assessing the impact of burning and grazing on the Konza Prairie Research Natural Area (a tallgrass prairie), spatial scale is extremely important. Textural contrast algorithms were applied to various scales of remote sensing data and related to landscape units for assessment of heterogeneity under a variety of burning treatments. Acquired data sets included Landsat multispectral scanner (MSS), with 80 m resolution, Landsat thematic mapper (TM), with 30 m resolution, and high resolution density sliced aerial photography (with a 5 m resolution). Results suggest that heterogeneous areas of dense patchiness (e.g., unburned areas) must be analyzed at a finer scale than more homogeneous areas which are burned at least every four years.     

Fire severity and seed source influence lodgepole pine (<Emphasis Type="Italic">Pinus contorta</Emphasis> var. <Emphasis Type="Italic">murrayana</Emphasis>) regeneration in the southern cascades,Lassen volcanic National Park,California

Rocky Mountain lodgepole pine, (Pinus contorta var. latifolia) regenerates quickly after high severity fire because seeds from serotinous cones are released immediately post-fire. Sierra lodgepole pine (P. contorta var. murrayana) forests burn with variable intensity resulting in different levels of severity and because this variety of lodgepole pine does not have serotinous cones, little is known about what factors influence post-fire regeneration. This study quantifies tree regeneration in a low, moderate, and high severity burn patch in a Sierra lodgepole forest 24 years after fire. Regeneration was measured in ten plots in each severity type. In each plot, we quantified pre- and post-fire forest structure (basal area, density), counted and aged tree seedlings and saplings of all species, and measured distance to the nearest seed bearing tree. There was no difference in the density of seedlings and saplings among severity classes. Distance and direction to the nearest seed bearing lodgepole pine were the best predictors of lodgepole seedling and sapling density in high severity plots. In contrast to Rocky Mountain lodgepole pine, regeneration of Sierra lodgepole pine appears to rely on in-seeding from surviving trees in low or moderate severity burn patches or live trees next to high severity burn patches. Our data demonstrate that Sierra lodgepole pine follows stand development pathways hypothesized for non-serotinous stands of Rocky Mountain lodgepole pine.     

Consequences of Landscape Heterogeneity on Grassland Diversity and Productivity in the Espinal Agroforestry System of Central Chile

The current land use system in the anthropogenic savannas (Espinales) of the Mediterranean climate region of Chile, has resulted in considerable heterogeneity at the landscape level which is associated with different covers of the legume tree, Acacia caven. The effects of landscape heterogeneity on the diversity and productivity of herbaceous plant communities were studied in 29 plots of 1000 m², with a wide range of woody cover. A detrended correspondence analysis of the species × plots matrix explained 73% of the total variation and revealed the existence of two trends of variation in floristic composition: one associated with physiographic position (hillsides and flatlands) and the other related to the number of years since the last cutting, or coppicing, of A. caven. Despite the great majority of the original herbaceous species having disappeared as a result of the prevailing land use system, some native species have been able to survive especially on hillside areas with low grazing intensity. Woody cover was a good indicator of spatial heterogeneity and land use history. It was also correlated with stocking rate, above-ground biomass of herbaceous vegetation, and soil fertility (organic matter, nitrogen and phosphorus concentration), both on hillsides and flatlands. The relationship between woody cover and herbaceous plant species richness was significant and unimodal in flat land areas, and linear, and marginally significant, on hillsides. The consequences of land use changes on the conservation of the ecological and productive values of grasslands are analyzed.     

Spatial variability in soil nitrogen dynamics after prescribed burning in Ohio mixed-oak forests

This study describes the results of the application of a single dormant season prescribed fire to two southern Ohio forest sites for the purposes of restoring the ecosystem functional properties that existed in these sites prior to major human intervention (clearcutting, fire suppression, and atmospheric deposition). Each forest site was composed of three contiguous watershed units, two of which were burned in April of 1996. The forest sites differed in soil pH and available litter mass prior to the fires, and in both sites pH and available inorganic N varied among landscape positions such that inorganic C increased with increasing longterm soil moisture potential (measured as the GIS-derived Integrated Moisture Index [IMI] developed for this region). The fire temperatures at 10 cm above the litter surface were generally 150–300°C, and 29–80% of the litter was consumed, depending on site and landscape position. Soil solution total inorganic N (TIN) present one month after the fires did not differ significantly from that present prior to the fires in either burned or unburned watersheds, but was consistently greater in mesic landscape positions than in more xeric ones. N mineralization potential and organic C content varied both among fires and landscape positions. At the site which burned at higher intensity, soil N mineralization and TIN were both decreased by fire. At the less intensely burned site, fire resulted in increased TIN in the soils from the more xeric landscape position, and greater soil organic C in soils from the intermediate soil moisture areas. Path analysis produced models for fire-induced changes in C and N dynamics capable of explaining 26–69% of the observed variation using combinations of landscape and fire behavior. Losses of N to volatilization from these single fires were generally < 1 kg N/ha, and thus could not be expected to ameliorate the effects of atmospheric N deposition in these sites.     

Behavior as a tool for assessing a managed landscape: a case study of the Karner blue butterfly

In an increasingly human-dominated landscape, effective management of disturbance-maintained ecosystems, such as grasslands and savannas, is critical to the conservation of biodiversity. Yet, the response of individual organisms to landscapes created by disturbances and management is rarely studied. In this study, we examined the endangered Karner blue butterfly, Lycaeides melissa samuelis, in a heterogeneous oak savanna. Our objective was to quantify the butterfly’s habitat use and behavior to assess the effects of prescribed burning. The oak savanna management in Ohio, USA divides each Karner blue site (n = 4) into three units. Each one-third unit is then burned, mowed, or unmanaged in an annual rotation within each site, and the result is a fire return interval of ~3 years. Our surveys measured habitat use, while behavior observations quantified reproduction and foraging for the two annual broods. Our habitat use results showed burned treatments were recolonized quickly, but there was not a clear selection for burned treatments. Foraging rates were similar in all treatments; however, females oviposited significantly less in unmanaged treatments (only 5 of 127 ovipositions). This oviposition preference was likely due to habitat degradation and the availability of recently burned, early successional habitat. Since Karner blues avoided reproduction in units unburned for ≥4 years, these units could be burned to create high quality early successional habitat. These results demonstrate how behavioral decisions can be pivotal forces driving spatial population dynamics. Our case study demonstrates how a fine-scale landscape perspective combined with measurements of behavioral processes can assist with management decision-making.     

Factors influencing female home range sizes in elk (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in North American landscapes

Home range size is a result of individual movements and the spatial distribution of a population. While body size, sex, and age are known to influence the area over which an animal ranges, it remains uncertain how landscape heterogeneity influences home range size. We examined elk (Cervus elaphus) seasonal home range sizes in relation to the quantity and spatial heterogeneity of forage biomass, forest cover, topography, snow–water equivalents, and landscape structure in three study landscapes: Yellowstone National Park, Wyoming, USA; eastern slopes of the Canadian Rockies, Alberta; and northern Wisconsin, USA. We used a 95% fixed kernel estimator to measure the home range size and location of all elk. To identify the scales at which important factors influenced home range sizes, we quantified environmental variables within the estimated home range polygon and within concentric circles with radii of 1000, 2000, 3000, 4000, and 5000 m from the home range center. Results indicate that there was an inverse relationship between forage biomass and summer and winter home range sizes in Alberta and Wisconsin, however the relationship was positive in Yellowstone. The size of summer and winter home ranges was positively related to percent forest cover; however this relationship was significant only when forest cover was quantified within the home range polygon or radii that were greater than or equal to 3000 m. Winter home ranges also had a positive relationship with snow–water equivalents. The predictive strength of summer home range models was greatest when landscape variables were quantified within the concentric circles with a radius of 3000 m or more, whereas the predictive strength of the winter models was greatest within the estimated home range polygon. Results suggest that elk ranging patterns reflected complex trade-offs that affect foraging, group dynamics, movement energetics, predation avoidance and thermal regulation. The multi-scale analysis indicates that elk based home ranging decisions on an area equal to their home range, but areas outside of the estimated home range were also important.     

Modelling the effects of landscape pattern and grazing regimes on the persistence of plant species with high conservation value in grasslands in south-eastern Sweden

Semi-natural grasslands in Sweden are threatened by land-use change and lack of management with attendant risk to their biodiversity. We present a model to explore the effects of grazing frequency and intensity on plant species persistence, and the relative effects of grassland size and pattern. We used a landscape modelling platform, LAMOS (LAndscape MOdelling Shell), to design a landscape model of vegetation dynamics incorporating the effects of local succession, dispersal and grazing disturbance. Five plant functional groups (PFG), representing various combinations of persistence and dispersal character, light requirements and disturbance responses, were defined to model species dynamics. Based on old cadastral maps three different landscapes were designed representing specific time-layers, i.e., a historical (17th to 18th century), a pre-modern (1940s) and a present-day landscape. Simulations showed that a threshold was crossed when grasslands decreased in area to about 10–30% of the modelled area, and as a consequence the biomass of grassland-specific PFGs was strongly reduced. These competition sensitive groups did not persist in the model even with intense grazing in the present-day landscape, where grasslands occupy 11% of the total area. However, all grassland species would have been able to persist in the historical landscape, where grasslands occupied 59% of the total area, even without grazing. Our results suggest that continuous but low-intensity grazing is more positive for grassland PFGs than discontinuous but highly intensive grazing. This effect was particularly strong when the frequency and/or intensity of grazing dropped below a threshold of 20%. Simulations using three landscape maps designed to explore effects of further fragmentation and habitat loss showed that the spatial pattern of remaining grasslands is important for the persistence of grassland-specific PFG. The model presented here is an advance towards more realistic grazing models to explore the effects of prescribed grazing and landscape fragmentation on the persistence species or plant functional groups.This revised version was published online in May 2005 with corrections to the Cover Date.     

Dispersion of kangaroo rat mounds at multiple scales in New Mexico,USA

Burrowing mammals create disturbances that increase the ecological heterogeneity of landscapes. In desert systems, banner-tailed kangaroo rats (Dipodomys spectabilis) construct large mounds that greatly influence the spatial patterning of soils, plants, and animals. The overall effects of the patches generated by D. spectabilis depend on the dispersion patterns of the mounds; these patterns may be sensitive to scale and landscape position. We examined the distribution of D. spectabilis mounds across multiple scales in four 40-ha grassland plots in New Mexico, USA. We used Ripley's K-function for our point-pattern analysis. The dispersion patterns of mounds were generally scale-sensitive but depended somewhat on plot-level densities, which were related to topographic position and grazing history. Mound spacing was either regular or random at small scales (0–50 m), random or aggregated at intermediate scales (50–300 m), and aggregated at large scales (300–3000 m). This scale-dependency of pattern reflected spatial domains in which different biotic (territoriality, dispersal, grazing) and abiotic (soil texture and drainage) factors exerted strong influences. How other organisms perceive the spatial patterning of mounds will depend on the extent of their movements. Patches may appear regular to one species but aggregated to another. The dispersion of D. spectabilis mounds also has implications for the spatial structuring of desert rodent communities. D. spectabilis excludes smaller species of kangaroo rats from areas around their mounds; they create spatial heterogeneity in behavioral dominance that may influence the distribution of subordinate species at multiple scales.     

Disturbances and gap dynamics in a semiarid grassland: A landscape-level approach

We developed a spatially-explicit gap dynamics simulation model to evaluate the effects of disturbances at the scale of a landscape for a semiarid grassland in northcentral Colorado, USA. The model simulates the establishment, growth, and death of individual plants on a small plot through time at an annual time step. Long-term successional dynamics on individual plots (single gaps) and on a landscape composed of a grid of plots were evaluated. Landscapes were simulated as either a collection of independent plots or as a collection of interacting plots where processes on one plot were influenced by processes on adjacent plots. Because we were interested in the recovery of the dominant plant species, the perennial grass blue grama (Bouteloua gracilis (H.B.K.) Lag. ex Griffiths) after disturbances, we focused on scale-dependent processes, such as seed dispersal, that are important to the recruitment of individuals of B. gracilis. The type of simulated landscape was important to the recovery time of B. gracilis after a disturbance. Landscapes composed of independent plots recovered more rapidly following a disturbance than landscapes composed of interacting plots in which the recovery time was dependent on the spatial scale of the disturbance.     

Landscape heterogeneity and fire behavior: scale-dependent feedback between fire and grazing processes

Fire and grazing are ecological processes that frequently interact to modify landscape patterns of vegetation. There is empirical and theoretical evidence that response of herbivores to heterogeneity is scale-dependent however the relationship between fire and scale of heterogeneity is not well defined. We examined the relationship between fire behavior and spatial scale (i.e., patch grain) of fuel heterogeneity. We created four heterogeneous landscapes modeled after those created by a fire–grazing interaction that differed in grain size of fuel patches. Fire spread was simulated through each model landscape from 80 independent, randomly located ignition points. Burn area, burn shape complexity and the proportion of area burnt by different fire types (headfire, backfire and flankfire) were all affected by the grain of fuel patch. The area fires burned in heterogeneous landscapes interacted with the fuel load present in the patch where ignition occurred. Burn complexity was greater in landscapes with small patch grain than in landscapes with large patch grain. The proportion of each fire type (backfire, flankfire and headfire) was similar among all landscapes regardless of patch grain but the variance of burned area within each of the three fire types differed among treatments of patch grain. Our landscape fire simulation supports the supposition that feedbacks between landscape patterns and ecological processes are scale-dependent, in this case spatial scale of fuel loading altering fire spread through the landscape.     

Quantifying the effect of grazing and shrub-clearing on small scale spatial pattern of vegetation

Disturbances such as grazing, invading species, and clear-cutting, often act at small spatial scales, and means for quantifying their impact on fine scale vegetation patterns are generally lacking. Here we adopt a set of landscape metrics, commonly used for quantifying coarse scale fragmentation, to quantify fine scale fragmentation, namely the fine scale vegetation structure. At this scale, patches often consist of individual plants smaller than 1 m², requiring the grain of the analysis to be much smaller. We used balloon aerial photographs to map fine details of Mediterranean vegetation (pixel size <0.04 m) in experimental plots subjected to grazing and clear-cutting and in undisturbed plots. Landscape metrics are sensitive to scale. Therefore, we aggregated the vegetation map into four coarser scales, up to a resolution of 1 m, and analyzed the effect of scale on the metrics and their ability to distinguish between different disturbances. At the finest scale, six of the seven landscape metrics we evaluated revealed significant differences between treated and undisturbed plots. Four metrics revealed differences between grazed and control plots, and six metrics revealed differences between cleared and control plots. The majority of metrics exhibited scaling relations. Aggregation had mixed effects on the differences between metric values for different disturbances. The control plots were the most sensitive to scale, followed by grazing and clearing. We conclude that landscape metrics are useful for quantifying the very fine scale impact of disturbance on woody vegetation, assuming that the analysis is based on sufficiently high spatial resolution data.     

Microhabitat conditions and landscape pattern explain nocturnal rodent activity,but not seed removal,in burned and unburned lodgepole pine forests

Context

Disturbances create spatial variation in environments that may influence animal foraging. Granivory by rodents can influence seed supply and thus plant establishment. However, effects of disturbance patterns on rodent seed removal in western North American conifer forests are generally unknown.

Objectives

We conducted a study in lodgepole pine (Pinus contorta var. latifolia) forests of Greater Yellowstone (Wyoming, USA) to answer: (1) How do seed removal and rodent activity vary between recently burned and adjacent unburned forests and with distance from fire perimeter? (2) Which microhabitat conditions explain variability in seed removal and rodent activity?

Methods

One or two years after wildfires, we established transects (n = 23) with four stations each: at 10 and 40 m from the fire perimeter in both burned and unburned forest. At stations, we deployed trays with lodgepole pine seeds and cameras pointed at trays for 28 days and quantified habitat structure and seed abundance.

Results

Seed removal, which averaged 85%, and diurnal rodent activity did not differ between burned and unburned forests or with distance from the fire perimeter; however, nocturnal rodent activity was lower in burned forests. Seed removal and diurnal rodent activity were not associated with any microhabitat conditions we measured. However, nocturnal rodent activity was associated with microhabitat in both burned and unburned forests.

Conclusions

High seed removal rates suggested that rodent foraging was not reduced by high-severity wildfire. If observed seed removal represents natural conditions, post-dispersal seed predation could influence post-fire recruitment of a widespread foundation tree species.

     

Wildfires and landscape patterns in the Eastern Iberian Peninsula

The relations between disturbance regime and landscape patterns have been developed from a theoretical perspective, but few studies have tested these relations when forces promoting opposing heterogeneity patterns are simultaneously operating on a landscape. This work provides quantitative evidence of these relations in areas dominated by human activity, showing that landscape heterogeneity decreases disturbance spread. In turn, disturbance introduces a source of landscape heterogeneity, but it is not enough to counterbalance the homogeneity trend due to agricultural abandonment. Land cover changes and wildfire occurrence (fires larger than 0.3 km²) have been monitored in the Tivissa municipality (208.4 km²) (Catalonia, NE Spain) from 1956 to 1993. Land cover maps were obtained from 1956, 1978 and 1993 and they were overlaid with fire occurrence maps obtained for the 1975–1995 period from 60 m resolution remote sensing images, which allow the identification of burned areas by sudden drops in Normalized Difference Vegetation Index (NDVI). Changes in landscape patterns in relation to fire regime have been analyzed considering several parameters: patch density, mean patch size, mean distance to the nearest neighbour of the same category, edge density, and the Shannon diversity index. In the 1956–1993 period there is a trend to increasing landscape homogenization due to the expansion of shrub­lands linked to a decrease in forest surface, and to the abandonment of agricultural lands. This trend, however, is not constant along all the period. Fires are more likely to occur in woody, homogenous areas, increasing landscape heterogeneity, as observed in the 1978–1993 period. This increase in heterogeneity does not counterbalance the general trend to landscape homogenization as a consequence of agricultural abandonment and the coalescence of natural vegetation patches.This revised version was published online in May 2005 with corrections to the Cover Date.     

Comparing the role of site disturbance and landscape properties on understory species richness in fragmented periurban Mediterranean forests

We hypothesized that the spatial configuration and dynamics of periurban forest patches in Barcelona (NE of Spain) played a minor role in determining plant species richness and assemblage compared to site conditions, and particularly to both direct (measured at plot level) and potential (inferred from landscape metrics) human-associated site disturbance. The presence of all understory vascular plants was recorded on 252 plots of 100 m² randomly selected within forest patches ranging in size from 0.25 ha to 218 ha. Species were divided into 6 groups, according to their ecology and conservation status. Site condition was assessed at plot level and included physical attributes, human-induced disturbance and Quercus spp. tree cover. Landscape structure and dynamics were assessed from patch metrics and patch history. We also calculated a set of landscape metrics related to potential human accessibility to forests. Results of multiple linear regressions indicated that the variance explained for non-forest species groups was higher than for forest species richness. Most of the main correlates corresponded to site disturbance variables related to direct human alteration, or to landscape variables associated to indirect human effects on forests: Quercus tree cover (a proxy for successional status) was the most important correlate of non-forest species richness, which decreased when Quercus tree cover increased. Human-induced disturbance was an important correlate of synanthropic and total species richness, which were higher in recently managed and in highly frequented forests. Potential human accessibility also affected the richness of most species groups. In contrast, patch size, patch shape and connectivity played a minor role, as did patch history. We conclude that human influence on species richness in periurban forests takes place on a small scale, whereas large-scale effects attributable to landscape structure and fragmentation are comparatively less important. Implications of these results for the conservation of plant species in periurban forests are discussed.     

Landscape context matters: local habitat and landscape effects on the abundance and patch occupancy of collared lizards in managed grasslands

The distribution and abundance of a species may be simultaneously influenced by both local-scale habitat features and the broader patch and landscape contexts in which these populations occur. Different factors may influence patch occupancy (presence–absence) versus local abundance (number of individuals within patches), and at different scales, and thus ideally both occupancy and abundance should be investigated, especially in studies that seek to understand the consequences of land management on species persistence. Our study evaluated the relative influences of variables associated with the local habitat patch, hillside (patch context), and landscape context on patch occupancy and abundance of the collared lizard (Crotaphytus collaris) within tallgrass prairie managed under different fire and grazing regimes in the northern Flint Hills of Kansas, USA. Using a multi-model information-theoretic approach that accounted for detection bias, we found that collared lizard abundance and occupancy was influenced by factors measured at both the local habitat and landscape scales. At a local scale, collared lizard abundance was greatest on large rock ledges that had lots of crevices, high vegetation complexity, and were located higher up on the hillslope. At the landscape scale, collared lizard abundance and occupancy were both higher in watersheds that were burned frequently (1–2 year intervals). Interestingly, grazing only had a significant effect on occupancy and abundance within less frequently burned (4-year burn interval) watersheds. Our results suggest that, in addition to the obvious habitat needs of this species (availability of suitable rock habitat), land-management practices have the potential to influence collared lizard presence and abundance in the grasslands of the Flint Hills. Thus, mapping the availability of suitable habitat is unlikely to be sufficient for evaluating species distributions and persistence in such cases without consideration of landscape management and disturbance history.     

Cost analysis of remotely sensed foraging paths in patchy landscapes with plant anti-herbivore defenses (Patagonia, Argentina)

We developed metrics at a landscape scale to evaluate the costs and rewards experienced by large herbivores while foraging in natural vegetation with patchy anti-herbivore plant structures. We show an application of these metrics to the analysis of 16,000 records of positions at successive 1 min intervals of free-ranging ewes (Ovis aries) harnessed with Global-Positioning-System (GPS) loggers, in a large paddock of the Patagonian Monte shrublands (Argentina). Dominant shrubs in the area display numerous anti-herbivore defenses (spiny-resinous leaves, thorny stems, etc.) protecting them from grazing and herbivore trampling. Preferred grasses and forbs constitute a minor part of aboveground plant biomass and grow in relatively open areas among or around shrub patches. We mapped the movement speed of ewes onto high-resolution aerial photographs of the grazed paddocks and estimated costs and rewards along their paths based on algorithms of surface cost theory. Ewes explored areas of sparse vegetation at low speeds compatible with predominant grazing, and increased their speed when crossing denser shrubby patches. The cost algorithm was applied to evaluate daily searching costs as well as grazing rewards in relation to the length of daily searching paths. The observed path lengths and search speeds were consistent with those that compensate costs and rewards of the grazing activities as estimated by the surface cost analysis. We conclude that the technique presented here constitutes a valuable tool to quantify the effect of landscape characteristics on behavioral traits of grazing animals in similar environments.     

Nutrient flows and land use change in a rural catchment: a modelling approach

Due largely to unprecedented land-use changes in the Porijõgi River catchment (southern Estonia) losses of nutrients and organic matter have decreased significantly. During the period 1987–1997 abandoned lands increased from 1.7 to 10.5% and arable lands decreased from 41.8 to 23.9%. At the same time, the runoff of total-N, total-P, SO₄ and organic matter (after BOD₅) decreased from 25.9 to 5.1, 0.32 to 0.13, 78 to 48, and 7.4 to 3.5 kg ha^–1 yr^–1, respectively. The most significant decreases occurred in agricultural subcatchments while the changes were insignificant in the forested upper course catchment. A simple empirical model which incorporates land-use pattern, fertilization intensity, soil parameters and water discharge accurately described the variations of total-N and total-P runoff in both the whole catchment and its agricultural subcatchments (R ² varies from 0.95–0.99 for N to 0.49–0.93 for P). In small agricultural subcatchments the rate of fertilization is found the most important factor for nitrogen runoff, whereas in larger mosaic watersheds land use pattern plays the main role. Seven alternative scenarios compiled on the base of the empirical model allow to forecast potential nitrogen and phosphorus losses from the catchment. This information can be used in further landscape and regional planning of the whole region.