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Fire ecology has a long history of empirical investigation in rangelands. However, the science is inconclusive and incomplete, sparking increasing interest on how to advance the discipline. Here, we introduce a new framework for qualitatively and quantitatively understanding the ranges of variability in fire regimes typical of experimental investigations in rangeland fire science compared with the range of conditions that actually occurred during contemporary social-ecological times. We implement this framework for one of rangelands’ most pyrogenic systems—the Great Plains of North America. We identify four social-ecological fire eras that have epitomized people’s relationship with wildland fire in the Great Plains since the last glacial maxima. These cultural fire eras include the now-extinct coexistence era (indigenous fire use), the suppression era (extermination of wildland fire occurrence), the shadow era (localized prescribed burning groups), and the emerging wildfire era (resulting from wildland fire management failures, continued decoupling of human-fire ignitions, and global change). Our synthesis demonstrates that experimental fire conditions have not explored the types and ranges of variation in fire regime components responsible for shaping rangeland vegetation—now, in the past, or into the future. Instead, scientific investigations have focused largely on controlling and minimizing sources of uncertainty and experimental variation, essentially eliminating ranges of variation that underpin the functioning of fire in modern social-ecological systems. Yet a series of scientific investigations exist that targeted a wider range of variability in fire regime components, leading to major advancements and the rejection of a number of long-standing rules of thumb in rangeland science and management. These include 1) the manipulation of fire return interval, 2) the pyric herbivory experiments, and 3) the extreme fire trials. We discuss the general philosophy shared among these studies, introduce scientific standards needed to avoid common pitfalls, and highlight opportunities to better understand how rangeland pattern and process correspond to critical ranges of variation in the human-fire relationship.

Quantifying resource selection (an organism's disproportionate use of available resources) is essential to infer habitat requirements of a species, develop management recommendations, predict species responses to changing conditions, and improve our understanding of the processes that underlie ecological patterns. Because study sites, even within the same region, can differ in both the amount and the arrangement of cover types, our objective was to determine whether proximal sites can yield markedly different resource selection results for a generalist bird, northern bobwhite (Colinus virginianus). We used 5 years of telemetry locations and newly developed land cover data at two, geographically distinct but relatively close sites in the south‐central semi‐arid prairies of North America. We fit a series of generalized linear mixed models and used an information‐theoretic model comparison approach to identify and compare resource selection patterns at each site. We determined that the importance of different cover types to northern bobwhite is site‐dependent on relatively similar and nearby sites. Specifically, whether bobwhite selected for shrub cover and whether they strongly avoided trees, depended on the study site in focus. Additionally, the spatial scale of selection was nearly an order of magnitude different between the cover types. Our study demonstrates that—even for one of the most intensively studied species in the world—we may oversimplify resource selection by using a single study site approach. Managing the trade‐offs between practical, generalized conclusions and precise but complex conclusions is one of the central challenges in applied ecology. However, we caution against setting recommendations for broad extents based on information gathered at small extents, even for a generalist species at adjacent sites. Before extrapolating information to areas beyond the data collected, managers should account for local differences in the availability, arrangement, and scaling of resources., Though it is known that animals can show conditional selection responses (e.g., differential habitat selection in different areas of their range, under variable weather conditions, or in response to variable food availability), we test a scenario where ecologists might reasonably assume habitat selection patterns to be static. Specifically, we found marked differences in the direction, magnitude, and scale of habitat selection by a generalist bird at two, proximal study sites in the same area of the species' range. Our study demonstrates that—even for one of the most intensively studied species in the world—we are likely to oversimplify habitat selection by using a single study site approach.

Human alteration of fire regimes is a hallmark of the Anthropocene; yet few studies have fully explored the implications of utilizing high- intensity fires in grasslands and savannas to manage shrub encroachment. Decades of fire research in South Africa inspired a unique convergence of high-intensity fire experiments in the USA. In the Great Plains of North America, high-intensity fire trials were designed to remove traditional investigator constraints that minimised variability in fire intensity and to explore woody mortality thresholds across a broader suite of experimental conditions. At the same time, studies in the Kruger National Park, South Africa, similarly investigated high-intensity fires to examine previously unstudied relationships between high-intensity fires and woody encroachment. These scientific pursuits have contributed to theoretical advances in our understanding of fire-vegetation dynamics. In this paper, we synthesise these high-intensity fire experiments, the empirical evidenceemerging from them and their importance for managing grassland and savanna ecosystems, and the lessons learned and challenges ahead to maintaining critical ranges of variation in fire regimes during the Anthropocene.&nbsp

On the Ground • We surveyed residents across Oklahoma about their awareness of prescribed fire. • Most respondents expressed support for prescribed fire for managing rangelands. • Although there was support for prescribed fire, few individuals implemented it. • Of the several reasons given for not burning, the most common were lack of training, lack of equipment, and choosing not to burn.

Widespread woody encroachment is a prominent concern for savanna systems as it is often accompanied by losses in productivity and biodiversity. Extensive ecosystem-level work has advanced our understanding of its causes and consequences. However, there is still debate over whether local management can override regional and global drivers of woody encroachment, and it remains largely unknown how encroachment influences woody community assemblages. Here, we examined species-level changes in woody plant distributions and size structure from the late 1980s to the late 2000s based on spatially intensive ground-based surveys across Kruger National Park, South Africa. This study region spans broad gradients in rainfall, soil texture, fire frequency, elephant density, and other topographic variables. Species-level changes in frequency of occurrence and size class proportion reflected widespread woody encroachment primarily by Dichrostachys cinerea and Combretum apiculatum, and a loss of large trees mostly of Sclerocarya birrea and Acacia nigrescens. Environmental variables determining woody species distributions across Kruger varied among species but did not change substantially between two sampling times, indicating that woody encroachers were thickening within their existing ranges. Overall, more areas across Kruger were found to have an increased number of common woody species through time, which indicated an increase in stem density. These areas were generally associated with decreasing fire frequency and rainfall but increasing elephant density. Our results suggest that woody encroachment is a widespread but highly variable trend across landscapes in Kruger National Park and potentially reflects an erosion of local heterogeneity in woody community assemblages. Many savanna managers, including in Kruger, aim to manage for heterogeneity in order to promote biodiversity, where homogenization of vegetation structure counters this specific goal. Increasing fire frequency has some potential as a local intervention. However, many common species increased in commonness even under near-constant disturbance conditions, which likely limits the potential for managing woody encroachment in the face of drivers beyond the scope of local control. Regular field sampling coupled with targeted fire management will enable more accurate monitoring of the rate of encroachment intensification.

A key pursuit in contemporary ecology is to differentiate regime shifts that are truly irreversible from those that are hysteretic. Many ecological regime shifts have been labeled as irreversible without exploring the full range of variability in stabilizing feedbacks that have the potential to drive an ecological regime shift back towards a desirable ecological regime. Removing fire from grasslands can drive a regime shift to juniper woodlands that cannot be reversed using typical fire frequency and intensity thresholds, and has thus been considered irreversible. This study uses a unique, long-term experimental fire landscape co-dominated by grassland and closed-canopy juniper woodland to determine whether extreme fire can shift a juniper woodland regime back to grassland dominance using aboveground herbaceous biomass as an indicator of regime identity. We use a space-for-time substitute to quantify herbaceous biomass following extreme fire in juniper woodland up to 15 years post-fire and compare these with (i) 15 years of adjacent grassland recovery post-fire, (ii) unburned closed-canopy juniper woodland reference sites and (iii) unburned grassland reference sites. Our results show grassland dominance rapidly emerges following fires that operate above typical fire intensity thresholds, indicating that grassland-juniper woodlands regimes are hysteretic rather than irreversible. One year following fire, total herbaceous biomass in burned juniper stands was comparable to grasslands sites, having increased from 5 ± 3 g m−2 to 142 ± 42 g m−2 (+2785 ± 812 percent). Herbaceous dominance in juniper stands continued to persist 15-years after initial treatment, reaching a maximum of 337 ± 42 g m−2 eight years post-fire. In juniper encroached grasslands, fires that operate above typical fire intensity thresholds can provide an effective method to reverse juniper woodland regime shifts. This has major implications for regions where juniper encroachment threatens rancher-based economies and grassland biodiversity and provides an example of how to operationalize resilience theory to disentangle irreversible thresholds from hysteretic system behavior.

Uncertainty in satellite-derived burned area estimates are especially high in grassland systems, which are some of the most frequently burned ecosystems in the world. In this study, we compare differences in predicted burned area estimates for a region with the highest fire activity in North America, the Flint Hills of Kansas, USA, using the moderate resolution imaging spectroradiometer (MODIS) MCD64A1 burned area product and a customization of the MODIS MCD64A1 product using a major ground-truthing effort by the Kansas Department of Health and Environment (KDHE-MODIS customization). Local-scale ground-truthing and the KDHE-MODIS product suggests MODIS burned area estimates under predicted fire occurrence by 28% over a 19-year period in the Flint Hills ecoregion. Between 2001 and 2019, MODIS product indicated 8% burned area for any given year in the Flint Hills, even in years when fire activity was highest (2008, 2009, 2011, 2014). Based on these results, coupling MODIS burned area computations with local scale ground-truth efforts has the potential to significantly improve fire occurrence estimates and reduce uncertainty in other grassland and savanna regions.

Woodland and agricultural expansion are major causes of grassland fragmentation. Fire and rainfall play important roles in maintaining grasslands, however, fire activity has been reduced in fragmented landscapes. Quantify the degree to which basic landscape fragmentation metrics could be used as drivers of woody cover potential. Woody plant percent cover was calculated between 2004 and 2008 at > 2000 sites. At each site, we calculated these fragmentation metrics for grassland cover type (classified by the National Land Cover Database); # patches, landscape proportion, edge density, largest patch index, effective mesh size and patch cohesion index within 3 circular areas (10 km2, 360 km2 and 3600 km2) surrounding the sampling site. A quantile regression was performed to identify which metrics were useful at predicting the 25th, 50th, 75th or 95th quantile of woody cover distribution. Grassland proportion and edge density were significant predictors of the woody plant potential (75th and 95th quantile). Woody cover potential was positively associated with edge density suggesting that fragmented areas (i.e., areas with high number of edges) maintained higher woody cover, while grassland proportion was negatively associated with woody plant potential. We propose that in addition to a lack of fire, fragmented landscapes may facilitate further woodland expansion by reducing natural land and restricting grasslands to smaller, less connected patches, which can maintain higher woody cover. Given current trends in woodland expansion, special attention should be given to areas that are found within a fragmented landscape and climatically prone to woodland expansion.

Woodland expansion is a global phenomenon that, despite receiving substantial attention in recent years, remains poorly understood. Landscape change of this magnitude has several impacts perceived as negative on landscape processes, such as influencing fire regimes, habitat for wildlife, and hydrological processes. In southern Great Plains, Juniperus virginiana has been identified as a major contributor to woodland expansion. Adding to the perplexity of this phenomenon is its evidence on numerous landscape types on several continents, documented under varying climates. Our study aimed to quantify a direct treatment to reduce or slow down woodland expansion in an experimental rangeland in central Oklahoma, United States under three treatments: 1) herbicide, 2) fire with herbicide, and 3) control (no fire, no herbicide) within areas classified as “open grassland” in 1979. Thereafter, we identified these same areas in 2010 with remotely sensed imagery (Light Detection And Ranging) to quantify 1) total encroachment and 2) total encroachment by three size classes: a) small 1 − 2.5 m, b) intermediate 2.5 − 4.5 m and c) tall > 4.5 m. Overall, of the total area classified as grassland in 1979 (277.64 ha), 31% had been encroached by 2010. Encroachment was greatest in the control treatments, followed by herbicide-only treatment application and lowest in the fire and herbicide treatment with minor differences in mean plant height (4.11 m ± 0.28). Encroached areas were mostly dominated by tall individuals (45 ± 3.5%), followed by the intermediate-height class (31.53 ± 1.10%) and the least recorded in the smallest-height class (23.46 ± 2.29%), suggesting expansion occurred during the initial phases of treatment application. The costly practice of herbicide application did not provide a feasible solution to control further woodland expansion. However, when using herbicide with fire, woodland expansion was reduced, highlighting the effectiveness of early intervention by fire in reducing encroachment. This further supports landscape-scale studies highlighting the effect of fire to reduce woodland expansion.

Grasslands are one of the most endangered ecosystems globally. Large tracts of grassland in the Great Plains, USA have been converted to cropland since the early 1900s, and woodlands are rapidly encroaching into many grasslands of the region due to changes in land management practices. Changes in the arrangement and proportion of different land cover types can affect biodiversity. We used bird survey data to identify the effect of land cover change on breeding bird ranges within the Great Plains over a 10-year period. Each species was categorized into one of the following habitat guilds: grassland, shrubland, woodland, wetland and generalist. We calculated the proportion of each land cover (including cropland, grassland, woodland, developed, barren, water bodies and wetlands) within a 1.6 km radius of each bird survey starting point. Within an occupancy modeling framework, we estimated colonization and extinction rates for each species and averaged them to the guild level. We also quantified changes in land cover from grassland to other cover types. Results show that grasslands were mostly converted to cropland and woodland, which were accompanied by positive extinction rates for certain grassland species. Extinction rates at the guild level were unrelated to increases in any land cover type, and observed land cover changes largely favored the shrubland guild. Overall, habitat-guild responses show little influence of woodland or cropland expansion at regional scales, although certain species ranges are predicted to decrease with increases in woodland. Future research should consider a finer scaled approach focusing on species-level responses when short-term land cover changes are considered.

Important drivers of woody cover in African savannas are rainfall, soil and nutrients, and disturbance factors, such as fire and herbivory. However, very little is known about how these drivers influence woody cover at specific height classes. The main aim of this study was to identify which of these drivers are the best predictors of woody cover at three height classes: shrub (0.75–2.5 m), brush (2.5–5.5 m) and tree (>5.5 m). Percent woody cover estimates were collected using a rapid monitoring technique over 1 700 sites in Kruger National Park, South Africa. Geology (basalt and granite), mean annual rainfall, fire frequency and elephant density were analysed as potential drivers of woody cover. Results indicate that mean annual rainfall was negatively associated with shrub cover, fire frequency was negatively associated with brush cover, and elephant density was negatively associated with tree cover. Patterns of woody cover show that while geology influences the spatial distribution of horizontal woody cover, variation in vertical vegetation structure is created and maintained by top-down disturbance. This provides the first documentation of drivers associated with structurally differentiated woody cover at regional scales. Future studies on woody cover in disturbance-driven environments should not ignore vegetation structure.Keywords: bush thickening, fire, herbivory, protected areas, woody vegetation

The present study was performed to determine whether the palatability, consumption and digestibility of an artificial sea urchin diet could be improved by the addition of a highly preferred seaweed. Four species of macroalgae (Ulva rigida, Ecklonia maxima, Porphyra capensis and Gigartina polycarpa) were assessed by pairwise preference tests. Each macroalgal species was consumed at least once by Tripneustes gratilla and significant preferences always involved Ulva as the most preferred species. Four protein-rich artificial diets supplemented with varying amounts of Ulva [0, 50, 150 and 200 g kg−1 designated 0, 5, 15 and 20U, respectively] were then developed and fed to urchins over a 20-day period. Inclusion of 200 g kg−1 dried Ulva significantly improved the palatability of the artificial diet and mean dry feed consumption rates were higher in urchins fed the 20U and 15U diets compared with urchins fed diets with a lower Ulva content. Daily digestible protein (DP) intake also differed significantly, with urchins fed the 20U diet having a significantly higher DP intake compared with the 15U, 5U and 0U treatments. These results indicate that inclusion of the palatable seaweed Ulva acted as a feeding stimulant, increasing the acceptability of the diets and boosting protein intake.

Nutrition is a major factor governing early development of sea urchins. The morphological development and survival of Tripneustes gratilla larvae, from field-collected urchins, was investigated from two separate feeding trials under different feeding regimes. Microalgae Isochrysis sp., Pavlova lutheri and Chaetoceros muelleri were tested as feed in May 2010 and Tetraselmis suecica, Skeletonema pseudocostatum and C. muelleri were tested in September 2010. Morphometric measurements of post-oral arm (PO), total width, total length, stomach length and mid-body line (MBL) were recorded every second day. In May, larvae fed Pavlova died after 23 days and did not reach competence. Larvae grew significantly better on Chaetoceros and Isochrysis, reaching competence on day 16 and 18 respectively, and had survival rates of 5.12% and 13.8% respectively. In the September trial, larvae fed Tetraselmis died by day 9, whereas larvae fed Skeletonema and Chaetoceros reached competence by day 22. Survival rates for Skeletone...

Woody plant cover and species composition play an important role in defining the type and function of savanna ecosystems. Approximately 2000 sites in the Kruger National Park (KNP) were surveyed by F.J. Venter over a period from 1985 to 1989, recording vegetation, soil and topological characteristics. At each of these sites (approximately 20 m × 20 m each), woody vegetation cover and species were recorded using a rapid, Braun-Blanquet classification for three height classes: shrub (0.75 m – 2.50 m), brush (2.50 m – 5.50 m) and tree (> 5.50 m). The objective of this study was to re-analyse the vegetation component of the field data, with a specific focus to provide a spatially explicit, height-differentiated, benchmark dataset in terms of species occurrence, species richness and structural canopy cover. Overall, 145 different woody species were recorded in the dataset out of the 458 species documented to occur in the park. The dataset describes a woody layer dominated by a relatively small number of widely occurring species, as 24 of the most common woody species accounted for all woody species found on over 80% of all sites. The less common woody species (101) were each recorded on 20 sites or less. Species richness varied from 12 to 1 species per site. Structural canopy cover averaged 9.34%, 8.16% and 2.89% for shrub, brush and tree cover, respectively. The dataset provides a useful benchmark for woody species distribution in KNP and can be used to explore woody species and height class distributions, as well as comparison with more recent or future woody vegetation surveys.Conservation implications: The results provided evidence that large-scale, woody vegetation surveys conducted along roads offer useful ecosystem level information. However, such an approach fails to pick up less common species. The data presented here provided a useful snapshot of KNP woody vegetation structure and composition and could provide excellent opportunities for spatio-temporal comparisons.

Woody plant cover and species composition play an important role in defining the type and function of savanna ecosystems. Approximately 2000 sites in the Kruger National Park (KNP) were surveyed by F.J. Venter over a period from 1985 to 1989, recording vegetation, soil and topological characteristics. At each of these sites (approximately 20 m × 20 m each), woody vegetation cover and species were recorded using a rapid, Braun-Blanquet classification for three height classes: shrub (0.75 m – 2.50 m), brush (2.50 m – 5.50 m) and tree (> 5.50 m). The objective of this study was to re-analyse the vegetation component of the field data, with a specific focus to provide a spatially explicit, height-differentiated, benchmark dataset in terms of species occurrence, species richness and structural canopy cover. Overall, 145 different woody species were recorded in the dataset out of the 458 species documented to occur in the park. The dataset describes a woody layer dominated by a relatively small number of widely occurring species, as 24 of the most common woody species accounted for all woody species found on over 80% of all sites. The less common woody species (101) were each recorded on 20 sites or less. Species richness varied from 12 to 1 species per site. Structural canopy cover averaged 9.34%, 8.16% and 2.89% for shrub, brush and tree cover, respectively. The dataset provides a useful benchmark for woody species distribution in KNP and can be used to explore woody species and height class distributions, as well as comparison with more recent or future woody vegetation surveys.Conservation implications: The results provided evidence that large-scale, woody vegetation surveys conducted along roads offer useful ecosystem level information. However, such an approach fails to pick up less common species. The data presented here provided a useful snapshot of KNP woody vegetation structure and composition and could provide excellent opportunities for spatio-temporal comparisons.

Understanding the dynamics of woody tree species distribution in savanna systems remains a challenge despite considerable attention the topic has received in recent years. Disturbances such as fire and elephant effects on woody vegetation are well documented, yet the influence of these factors on emerging landscape-scale patterns such as height and species distributions continue to be poorly understood. The aim of this study was to identify how a suite of environmental variables (rainfall, temperature, aspect, slope, geology, fire frequency and elephant density) and their relative contributions, may affect woody species distribution in relation to structural height classes. Using the Maximum Entropy model for three structural height classes of the fifteen most frequently occurring woody species in the Kruger National Park (South Africa), the environmental variables best explaining each species distribution were identified. The three structural classes were defined to capture canopy height categories of sp...