Should heterogeneity be the basis for conservation? Grassland bird response to fire and grazing.

In tallgrass prairie, disturbances such as grazing and fire can generate patchiness across the landscape, contributing to a shifting mosaic that presumably enhances biodiversity. Grassland birds evolved within the context of this shifting mosaic, with some species restricted to one or two patch types created under spatially and temporally distinct disturbance regimes. Thus, management-driven reductions in heterogeneity may be partly responsible for declines in numbers of grassland birds. We experimentally altered spatial heterogeneity of vegetation structure within a tallgrass prairie by varying the spatial and temporal extent of fire and by allowing grazing animals to move freely among burned and unburned patches (patch treatment). We contrasted this disturbance regime with traditional agricultural management of the region that promotes homogeneity (traditional treatment). We monitored grassland bird abundance during the breeding seasons of 2001-2003 to determine the influence of altered spatial heterogeneity on the grassland bird community. Focal disturbances of patch burning and grazing that shifted through the landscape over several years resulted in a more heterogeneous pattern of vegetation than uniform application of fire and grazing. Greater spatial heterogeneity in vegetation provided greater variability in the grassland bird community. Some bird species occurred in greatest abundance within focally disturbed patches, while others occurred in relatively undisturbed patches in our patch treatment. Henslow's Sparrow, a declining species, occurred only within the patch treatment. Upland Sandpiper and some other species were more abundant on recently disturbed patches within the same treatment. The patch burn treatment created the entire gradient of vegetation structure required to maintain a suite of grassland bird species that differ in habitat preferences. Our study demonstrated that increasing spatial and temporal heterogeneity of disturbance in grasslands increases variability in vegetation structure that results in greater variability at higher trophic levels. Thus, management that creates a shifting mosaic using spatially and temporally discrete disturbances in grasslands can be a useful tool in conservation. In the case of North American tallgrass prairie, discrete fires that capitalize on preferential grazing behavior of large ungulates promote a shifting mosaic of habitat types that maintain biodiversity and agricultural productivity.     

Pyric Herbivory: Rewilding Landscapes through the Recoupling of Fire and Grazing

Abstract:  Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison ( Bison bison ) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances .     

Weak climatic control of stand-scale fire history during the late holocene

Forest fire occurrence is affected by multiple controls that operate at local to regional scales. At the spatial scale of forest stands, regional climatic controls may be obscured by local controls (e.g., stochastic ignitions, topography, and fuel loads), but the long-term role of such local controls is poorly understood. We report here stand-scale (<100 ha) fire histories of the past 5000 years based on the analysis of sediment charcoal at two lakes 11 km apart in southeastern British Columbia. The two lakes are today located in similar subalpine forests, and they likely have experienced the same late-Holocene climatic changes because of their close proximity. We evaluated two independent properties of fire history: (1) fire-interval distribution, a measure of the overall incidence of fire, and (2) fire synchroneity, a measure of the co-occurrence of fire (here, assessed at centennial to millennial time scales due to the resolution of sediment records). Fire-interval distributions differed between the sites prior to, but not after, 2500 yr before present. When the entire 5000-yr period is considered, no statistical synchrony between fire-episode dates existed between the two sites at any temporal scale, but for the last 2500 yr marginal levels of synchrony occurred at centennial scales. Each individual fire record exhibited little coherency with regional climate changes. In contrast, variations in the composite record (average of both sites) matched variations in climate evidenced by late-Holocene glacial advances. This was probably due to the increased sample size and spatial extent represented by the composite record (up to 200 ha) plus increased regional climatic variability over the last several millennia, which may have partially overridden local, non-climatic controls. We conclude that (1) over past millennia, neighboring stands with similar modern conditions may have experienced different fire intervals and asynchronous patterns in fire episodes, likely because local controls outweighed the synchronizing effect of climate; (2) the influence of climate on fire occurrence is more strongly expressed when climatic variability is relatively great; and (3) multiple records from a region are essential if climate-fire relations are to be reliably described.     

Effect of sheep grazing and fire on sage grouse populations in southeastern Idaho

This paper describes the development, evaluation, and use of a model that simulates the effect of grazing and fire on temporal and spatial aspects of sagebrush community vegetation and sage grouse population dynamics. The model is represented mathematically as a discrete-time, stochastic compartment model based on difference equations with a time interval of 1 week. In the model, sheep graze through sage grouse breeding habitat during spring and fall, and different portions of the area can burn at different frequencies, creating a habitat mosaic of burned and unburned areas.The model was evaluated by examining predictions of (1) growth of sagebrush canopy cover after fire, (2) seasonal dynamics of grass and forb biomass under historical environmental conditions, and (3) sage grouse population dynamics associated with selected sagebrush canopy covers. Simulated changes in sagebrush canopy cover following fire correspond well with qualitative reports of long-term trends, simulated seasonal dynamics of herbaceous biomass correspond well with field data, and simulated responses of sage grouse population size and age structure to changing sagebrush canopy cover correspond well to qualitative field observations.Simulation results suggest that large fires occurring at high frequencies may lead to the extinction of sage grouse populations, whereas fires occurring at low frequencies may benefit sage grouse if burned areas are small and sheep grazing is absent. Sheep grazing may contribute to sage grouse population decline, but is unlikely to cause extinction under fire regimes that are favorable to sage grouse.     

Postfire response of flood-regenerating riparian vegetation in a semi-arid landscape

Piles of large wood (LW) deposited by major floods in river corridors can interact with naturally occurring wildfires from uplands to impact the regeneration of riparian vegetation. This study examines the spatial and short-term temporal response of riparian vegetation and soil nutrients to fire along the Sabie River, South Africa, with special emphasis on the effects of burned LW piles. At the study site there were 112 species of plants recorded with 28% of species restricted to the burned plots. As expected, vegetation cover was significantly lower in burned plots as compared with the unburned plots 12 months postfire. There was a significant influence of LW on species richness with fewer species recorded in the LW plots. For both fire and LW treatments, plant cover showed a significant change over three years. After an initial increase from 12 to 24 months (postfire) there was a decline in plant cover after 36 months. Species community composition was distinctly different between burned and unburned plots 12 months postfire, and the presence of LW affected species composition for burned plots but not for unburned ones. Time series ordination of LW plots highlighted the changes in species composition over the three years of sampling. Of trees with accumulations of LW within 5 m of their base, 48% had been killed by fire as compared to only 4% with no LW accumulations in close proximity. Soil-available P was significantly higher in the burned plots and even higher with burned LW while there were no effects on soil total N. There was also a significant positive trend between available P in soils and plant vegetation cover. Soil-exchangeable K was also significantly higher and total C significantly lower in the burned and LW plots. Burned plots also had significantly higher soil electrical conductivity (EC) and soil pH. The patchy nature of the studied fire, whose complexity is exacerbated by the distribution of flood deposited LW, acted to create a mosaic of alternate successional states as the riparian community recovers from flooding and the subsequent fire. We suspect that the resultant heterogeneity will increase ecosystem resilience by providing flexibility in the form of more options for a system response to subsequent disturbances.     

Scale-dependent variation in coral community similarity across sites, islands, and island groups

Community similarity is the proportion of species richness in a region that is shared on average among communities within that region. The slope of local richness (alpha diversity) regressed on regional richness (gamma diversity) can serve as an index of community similarity across regions with different regional richness. We examined community similarity in corals at three spatial scales (among transects at a site, sites on an island, and islands within an island group) across a 10 000-km longitudinal diversity gradient in the west-central Pacific Ocean. When alpha diversity was regressed on gamma diversity, the slopes, and thus community similarity, increased with scale (0.085, 0.261, and 0.407, respectively) because a greater proportion of gamma diversity was subsumed within alpha diversity as scale increased. Using standard randomization methods, we also examined how community similarity differed between observed and randomized assemblages and how this difference was affected by spatial separation of species within habitat types and specialization of species to three habitat types (reef flats, crests, and slopes). If spatial separation within habitat types and/or habitat specialization (i.e., underdispersion) occurs, fewer species are shared among assemblages than the random expectation. When the locations of individual coral colonies were randomized within and among habitat types, community similarity was 46-47% higher than that for observed assemblages at all three scales. We predicted that spatial separation of coral species within habitat types should increase with scale due to dispersal/extinction dynamics in this insular system, but that specialization of species to different habitat types should not change because habitat differences do not change with scale. However, neither habitat specialization nor spatial separation within habitat types differed among scales. At the two larger scales, each accounted for 22-24% of the difference in community similarity between observed and randomized assemblages. At the smallest scale (transect-site), neither spatial separation within habitat types nor habitat specialization had significant effects on community similarity, probably due to the small size of transect samples. The results suggest that coral species can disperse among islands in an island group as easily as they can among sites on an island over time scales that are relevant to their establishment and persistence on reefs.     

Community occupancy responses of small mammals to restoration treatments in ponderosa pine forests, northern Arizona, USA

In western North American conifer forests, wildfires are increasing in frequency and severity due to heavy fuel loads that have accumulated after a century of fire suppression. Forest restoration treatments (e.g., thinning and/or burning) are being designed and implemented at large spatial and temporal scales in an effort to reduce fire risk and restore forest structure and function. In ponderosa pine (Pinus ponderosa) forests, predominantly open forest structure and a frequent, low-severity fire regime constituted the evolutionary environment for wildlife that persisted for thousands of years. Small mammals are important in forest ecosystems as prey and in affecting primary production and decomposition. During 2006-2009, we trapped eight species of small mammals at 294 sites in northern Arizona and used occupancy modeling to determine community responses to thinning and habitat features. The most important covariates in predicting small mammal occupancy were understory vegetation cover, large snags, and treatment. Our analysis identified two generalist species found at relatively high occupancy rates across all sites, four open-forest species that responded positively to treatment, and two dense-forest species that responded negatively to treatment unless specific habitat features were retained. Our results indicate that all eight small mammal species can benefit from restoration treatments, particularly if aspects of their evolutionary environment (e.g., large trees, snags, woody debris) are restored. The occupancy modeling approach we used resulted in precise species-level estimates of occupancy in response to habitat attributes for a greater number of small mammal species than in other comparable studies. We recommend our approach for other studies faced with high variability and broad spatial and temporal scales in assessing impacts of treatments or habitat alteration on wildlife species. Moreover, since forest planning efforts are increasingly focusing on progressively larger treatment implementation, better and more efficiently obtained ecological information is needed to inform these efforts.     

Beta diversity at different spatial scales: plant communities in organic and conventional agriculture.

Biodiversity studies that guide agricultural subsidy policy have generally compared farming systems at a single spatial scale: the field. However, diversity patterns vary across spatial scales. Here, we examined the effects of farming system (organic vs. conventional) and position in the field (edge vs. center) on plant species richness in wheat fields at three spatial scales. We quantified alpha-, beta-, and gamma-diversity at the microscale in 800 plots, at the mesoscale in 40 fields, and at the macroscale in three regions using the additive partitioning approach, and evaluated the relative contribution of beta-diversity at each spatial scale to total observed species richness. We found that alpha-, beta-, and gamma-diversity were higher in organic than conventional fields and higher at the field edge than in the field center at all spatial scales. In both farming systems, beta-diversity at the meso- and macroscale explained most of the overall species richness (up to 37% and 25%, respectively), indicating considerable differences in community composition among fields and regions due to environmental heterogeneity. The spatial scale at which beta-diversity contributed the most to overall species richness differed between rare and common species. Total richness of rare species (present in < or = 5% of total samples) was mainly explained by differences in community composition at the meso- and macroscale (up to 27% and 48%, respectively), but only in organic fields. Total richness of common species (present in > or = 25% of total samples) was explained by differences in community composition at the micro- and mesoscale (up to 29% and 47%, respectively), i.e., among plots and fields, independent of farming system. Our results show that organic farming made the greatest contribution to total species richness at the meso (among fields) and macro (among regions) scale due to environmental heterogeneity. Hence, agri-environment schemes should exploit this large-scale contribution of beta-diversity by tailoring schemes at regional scales to maximize dissimilarity between conservation areas using geographic information systems rather than focusing entirely at the classical local-field scale, which is the current practice.     

Fire severity and ecosytem responses following crown fires in California shrublands

Chaparral shrublands burn in large high-intensity crown fires. Managers interested in how these wildfires affect ecosystem processes generally rely on surrogate measures of fire intensity known as fire severity metrics. In shrublands burned in the autumn of 2003, a study of 250 sites investigated factors determining fire severity and ecosystem responses. Using structural equation modeling we show that stand age, prefire shrub density, and the shortest interval of the prior fire history had significant direct effects on fire severity, explaining > 50% of the variation in severity. Fire severity per se is of interest to resource managers primarily because it is presumed to be an indicator of important ecosystem processes such as vegetative regeneration, community recovery, and erosion. Fire severity contributed relatively little to explaining patterns of regeneration after fire. Two generalizations can be drawn: fire severity effects are mostly shortlived, i.e., by the second year they are greatly diminished, and fire severity may have opposite effects on different functional types. Species richness exhibited a negative relationship to fire severity in the first year, but fire severity impacts were substantially less in the second postfire year and varied by functional type. Much of this relationship was due to alien plants that are sensitive to high fire severity; at all scales from 1 to 1000 m2, the percentage of alien species in the postfire flora declined with increased fire severity. Other aspects of disturbance history are also important determinants of alien cover and richness as both increased with the number of times the site had burned and decreased with time since last fire. A substantial number of studies have shown that remote-sensing indices are correlated with field measurements of fire severity. Across our sites, absolute differenced normalized burn ratio (dNBR) was strongly correlated with field measures of fire severity and with fire history at a site but relative dNBR was not. Despite being correlated with fire severity, absolute dNBR showed little or no relationship with important ecosystem responses to wildfire such as shrub resprouting or total vegetative regeneration. These findings point to a critical need for further research on interpreting remote sensing indices as applied to postfire management of these shrublands.     

Fire Regime Changes in La Michilía Biosphere Reserve, Durango, Mexico

Abstract: The ability of reserves to maintain natural ecosystem processes such as fire disturbance regimes is central to long-term conservation. Fire-scarred tree samples were used to reconstruct fire regimes at five study sites totaling approximately 230 ha in pine (   Pinus spp.) and oak ( Quercus spp.) forests of La Michilía Biosphere Reserve on the dry east slope of the Sierra Madre Occidental, Durango, Mexico. Study sites covered a 20-km environmental gradient of elevation, topography, and human land uses. Plant communities ranged from oak-pine to mixed conifer forests. Fires were frequent at all sites prior to 1930, when large-scale grazing of domestic livestock was initiated. Widespread fires have been excluded from four of the five sites since 1945, with an essentially uninterrupted regime of frequent fires continuing only in the reserve core. Xeric sites had many, smaller fires, whereas mesic sites had fewer but larger fires. On a reserve-wide scale, a fire burned on at least one site nearly every year, usually in the dry spring or early summer season, but fire years were rarely synchronous among the sites. Fire occurrence was weakly related to the Southern Oscillation climate pattern; major reserve-wide fire years almost never coincided with wet Southern Oscillation extremes but only occasionally matched dry extremes. Maintenance of the long-term frequent-fire regime in the reserve core is one indicator that the biosphere reserve model has been successful in conserving natural processes, but the protected area is small ( 7000 ha). Because of the key role of frequent-fire regimes in regulating ecosystem structure and function, restoration of the ecological role of fire disturbance is a desirable conservation strategy.     

Small-scale fuel variation alters fire intensity and shrub abundance in a pine savanna

Small-scale variation in fire intensity and effects may be an important source of environmental heterogeneity in frequently burned plant communities. We hypothesized that variation in fire intensity resulting from local differences in fuel loads produces heterogeneity in pine savanna ground cover by altering shrub abundance. To test this hypothesis, we experimentally manipulated prefire fuel loads to mimic naturally occurring fuel-load heterogeneity associated with branch falls, needle fall near large pines, and animal disturbances in a frequently burned longleaf pine (Pinus palustris) savanna in Louisiana, USA. We applied one of four fuel treatments (unaltered control, fine-fuel removal, fine-fuel addition, wood addition) to each of 540 (1-m2) quadrats prior to growing-season prescribed fires in each of two years (1999 and 2001). In both years fuel addition increased (and fuel removal decreased) fuel consumption and maximum fire temperatures relative to unaltered controls. Fuel addition, particularly wood, increased damage to shrubs, increased shrub mortality, and decreased resprout density relative to controls. We propose that local variation in fire intensity may contribute to maintenance of high species diversity in pine savannas by reducing shrub abundance and creating openings in an otherwise continuous ground cover.     

The influence of aridity and fire on holocene prairie communities in the eastern Prairie Peninsula

The role of climate and fire in the development, maintenance, and species composition of prairie in the eastern axis of the tallgrass Prairie Peninsula intrigued early North American ecologists. However, evaluation of the long-standing hypotheses about the region's environmental history has been hampered by the scarcity of paleorecords. We conducted multiproxy analyses on early and middle Holocene sediments from two Illinois, USA, lakes to assess long-term climatic, vegetational, and fire variability in the region. Sediment mineral composition, carbonate delta18O, ostracode assemblages, and diatom assemblages were integrated to infer fluctuations in moisture availability. Pollen and charcoal delta13C were used to reconstruct vegetation composition, and charcoal influx was used to reconstruct fire. Results indicate that fire-sensitive trees (e.g., Ulmus, Ostrya, Fraxinus, and Acer saccharum) declined and prairie taxa expanded with increased aridity from 10,000 yr BP to 8500 yr BP. Between approximately 8500 yr BP and approximately 6200 yr BP, aridity declined, and prairie coexisted with fire-sensitive and fire-tolerant (e.g., Quercus and Carya) trees. After approximately 6200 yr BP, prairie taxa became dominant, although aridity was not more severe than it was around 8500 yr BP. Along with aridity, fire appears to have played an important role in the establishment and maintenance of prairie communities in the eastern Prairie Peninsula, consistent with the speculations of the early ecologists. Comparison of our data with results from elsewhere in the North American midcontinent indicates that spatial heterogeneity is a characteristic feature of climatic and vegetational variations on millennial time scales.     

Dispersal limitation and environmental heterogeneity shape scale-dependent diversity patterns in plant communities

Understanding the large-scale distribution of species diversity requires distinguishing two of the primary factors that cause compositional differences: dispersal limitation and environmental variation. In a community with a naturally discontinuous spatial structure, we asked (1) at what scale(s) nonrandom variation in species composition occurs and (2) at what scale(s) such variation is associated with spatial separation, indicative of dispersal limitation, and at what scale(s) variation is associated with environmental heterogeneity? We sampled 50 seeps (small wetlands) on five serpentine outcrops. Using a randomization model, we showed that additive beta diversity (a measure of community dissimilarity) was lower than random within seeps and higher than random among both seeps and outcrops. Using Mantel tests, we showed that plant community dissimilarity, in both the full seep assemblage as well as in a subset of seep endemics, at the two larger scales was associated with different forms of environmental heterogeneity and, at the largest scale, was also associated with geographic distance. We conclude that diversity in this system is shaped by multiple scales of heterogeneity and by dispersal limitation at the largest scale.     

Landsat Thematic Mapper: Detection of Shifts in Community Composition of Coral Reefs

Abstract: We assembled a time series of 20 Landsat thematic mapper images from 1982 to 1996 for Key Largo, Florida, to ascertain whether satellite imagery can detect temporal changes in coral reef communities. Selected reef and control areas were examined for changes in brightness, spectral reflectance, band ratios, spatial texture, and temporal texture (  pixel-to-pixel change over time). We compared the data to known changes in the reef ecosystem of Carysfort Reef and terrestrial sample sites. Changes in image brightness and spectral-band ratios were suggestive of shifts from coral- to algal-dominated community structure, but the trends were not statistically significant. The spatial heterogeneity of the reef community decreased in the early 1980s at scales consistent with known ecological changes to the coral community on Carysfort Reef. An analysis of pixel-scale variation through time, termed temporal texture, revealed that the shallow reef areas are the most variable in regions of the reef that have experienced significant ecological decline. Thus, the process of reef degradation, which alters both the spatial patterning and variability of pixel brightness, can be identified in unclassified thematic mapper images.     

Application of the New Keystone-Species Concept to Prairie Dogs: How Well Does It Work?

Abstract: It has been suggested that the keystone-species concept should be dropped from ecology and conservation, primarily because the concept is poorly defined. This prompted Power et al. (1996) to refine the definition: keystone species have large effects on community structure or ecosystem function (i.e., high overall importance), and this effect should be large relative to abundance (i.e., high community importance). Using prairie dogs ( Cynomys spp.) as an example, I review operational and conceptual difficulties encountered in applying this definition. As applied to prairie dogs, the implicit assumption that overall importance is a linear function of abundance is invalid. In addition, community importance is sensitive to abundance levels, the definition of community, and sampling scale. These problems arise largely from the equation for community importance, as used in conjunction with removal experiments at single abundance levels. I suggest that we shift from the current emphasis on the dualism between keystone and nonkeystone species and instead examine how overall and community importance vary (1) with abundance, (2) across spatial and temporal scales, and (3) under diverse ecological conditions. In addition, I propose that a third criterion be incorporated into the definition: keystone species perform roles not performed by other species or processes. Examination of how these factors vary among populations of keystone species should help identify the factors contributing to, or limiting, keystone-level functions, thereby increasing the usefulness of the keystone-species concept in ecology and conservation. Although the quantitative framework of Power et al. falls short of being fully operational, my conceptual guidelines may improve the usefulness of the keystone-species concept. Careful attention to the factors that limit keystone function will help avoid misplaced emphasis on keystone species at the expense of other species.     

Trends and causes of severity, size, and number of fires in northwestern California, USA

Research in the last several years has indicated that fire size and frequency are on the rise in western U.S. forests. Although fire size and frequency are important, they do not necessarily scale with ecosystem effects of fire, as different ecosystems have different ecological and evolutionary relationships with fire. Our study assessed trends and patterns in fire size and frequency from 1910 to 2008 (all fires > 40 ha), and the percentage of high-severity in fires from 1987 to 2008 (all fires > 400 ha) on the four national forests of northwestern California. During 1910-2008, mean and maximum fire size and total annual area burned increased, but we found no temporal trend in the percentage of high-severity fire during 1987-2008. The time series of severity data was strongly influenced by four years with region-wide lightning events that burned huge areas at primarily low-moderate severity. Regional fire rotation reached a high of 974 years in 1984 and fell to 95 years by 2008. The percentage of high-severity fire in conifer-dominated forests was generally higher in areas dominated by smaller-diameter trees than in areas with larger-diameter trees. For Douglas-fir forests, the percentage of high-severity fire did not differ significantly between areas that re-burned and areas that only burned once (10% vs. 9%) when re-burned within 30 years. Percentage of high-severity fire decreased to 5% when intervals between first and second fires were > 30 years. In contrast, in both mixed-conifer and fir/high-elevation conifer forests, the percentage of high-severity fire was less when re-burned within 30 years compared to first-time burned (12% vs. 16% for mixed conifer; 11% vs. 19% for fir/high-elevation conifer). Additionally, the percentage of high-severity fire did not differ whether the re-burn interval was less than or greater than 30 years. Years with larger fires and greatest area burned were produced by region-wide lightning events, and characterized by less winter and spring precipitation than years dominated by smaller human-ignited fires. Overall percentage of high-severity fire was generally less in years characterized by these region-wide lightning events. Our results suggest that, under certain conditions, wildfires could be more extensively used to achieve ecological and management objectives in northwestern California.     

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.     

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.     

Response of Savanna Fire Regimes to Changing Fire-Management Policies in a Large African National Park

Abstract:  Approaches to fire management in the savanna ecosystems of the 2-million ha Kruger National Park, South Africa, have changed several times over the past six decades. These approaches have included regular and flexible prescribed burning on fixed areas and a policy that sought to establish a lightning-dominated fire regime. We sought to establish whether changes in management induced the desired variability in fire regimes over a large area. We used a spatial database of information on all fires in the park between 1957 and 2002 to determine elements of the fire regime associated with each management policy. The area that burned in any given year was independent of the management approach and was strongly related to rainfall (and therefore grass fuels) in the preceding 2 years. On the other hand, management did affect the spatial heterogeneity of fires and their seasonal distribution. Heterogeneity was higher at all scales during the era of prescribed burning, compared with the lightning-fire interval. The lightning-fire interval also resulted in a greater proportion (72% vs. 38%) of the area burning in the dry season. Mean fire-return intervals varied between 5.6 and 7.3 years, and variability in fire-return intervals was strongly influenced by the sequencing of annual rainfall rather than by management. The attempt at creating a lightning-dominated fire regime failed because most fires were ignited by humans, and the policy has been replaced by a more pragmatic approach that combines flexible prescribed burning with lightning-ignited fires.     

Associations of Grassland Bird Communities with Black‐Tailed Prairie Dogs in the North American Great Plains

Colonial burrowing herbivores can modify vegetation structure, create belowground refugia, and generate landscape heterogeneity, thereby affecting the distribution and abundance of associated species. Black‐tailed prairie dogs (Cynomys ludovicianus) are such a species, and they may strongly affect the abundance and composition of grassland bird communities. We examined how prairie dog colonies in the North American Great Plains affect bird species and community composition. Areas occupied by prairie dogs, characterized by low percent cover of grass, high percent cover of bare soil, and low vegetation height and density, supported a breeding bird community that differed substantially from surrounding areas that lacked prairie dogs. Bird communities on colony sites had significantly greater densities of large‐bodied carnivores (Burrowing Owls [Athene cunicularia], Mountain Plovers, [Charadrius montanus], and Killdeer [Charadrius vociferus]) and omnivores consisting of Horned Larks (Eremophila alpestris) and McCown's Longspurs (Rhynchophanes mccownii) than bird communities off colony sites. Bird communities off colony sites were dominated by small‐bodied insectivorous sparrows (Ammodramus spp.) and omnivorous Lark Buntings (Calamospiza melanocorys), Vesper Sparrows (Pooecetes gramineus), and Lark Sparrows (Chondestes grammacus). Densities of 3 species of conservation concern and 1 game species were significantly higher on colony sites than off colony sites, and the strength of prairie dog effects was consistent across the northern Great Plains. Vegetation modification by prairie dogs sustains a diverse suite of bird species in these grasslands. Collectively, our findings and those from previous studies show that areas in the North American Great Plains with prairie dog colonies support higher densities of at least 9 vertebrate species than sites without colonies. Prairie dogs affect habitat for these species through multiple pathways, including creation of belowground refugia, supply of prey for specialized predators, modification of vegetation structure within colonies, and increased landscape heterogeneity. Asociaciones de Comunidades de Aves de Pastizales con Perros de la Pradera en la Gran Llanura de Norte América