Climate change causes declines and greater extremes in wetland inundation in a region important for wetland birds.

Wetland ecosystems are vital for maintaining global biodiversity, as they provide important stopover sites for many species of migrating wetland-associated birds. However, because weather determines their hydrologic cycles, wetlands are highly vulnerable to effects of climate change. Although changes in temperature and precipitation resulting from climate change are expected to reduce inundation of wetlands, few efforts have been made to quantify how these changes will influence the availability of stopover sites for migratory wetland birds. Additionally, few studies have evaluated how climate change will influence interannual variability or the frequency of extremes in wetland availability. For spring and fall bird migration in seven ecoregions in the south-central Great Plains of North America, we developed predictive models associating abundance of inundated wetlands with a suite of weather and land cover variables. We then used these models to generate predictions of wetland inundation at the end of the century (2069-2099) under future climate change scenarios. Climate models predicted the average number of inundated wetlands will likely decline during both spring and fall migration periods, with declines being greatest in the eastern ecoregions of the southern Great Plains. However, the magnitude of predicted declines varied considerably across climate models and ecoregions, with uncertainty among climate models being greatest in the High Plains ecoregion. Most ecoregions also were predicted to experience more-frequent extremely dry years (i.e., years with extremely low wetland abundances), but the projected change in interannual variability of wetland inundation was relatively small and varied across ecoregions and seasons. Because the south-central Great Plains represents an important link along the migratory routes of many wetland-dependent avian species, future declines in wetland inundation and more frequent periods of only a few wetlands being inundated will result in an uncertain future for migratory birds as they experience reduced availability of wetland stopover habitat across their migration pathways.

Climate Change and Wetlands in the Southern Great Plains: How Are Managers Dealing with an Uncertain Future?

Little guidance is available to assist wetland managers in developing climate adaptation plans. To facilitate development of recommendations for adaptation strategies, it is essential to first determine if or how wetland managers are addressing these challenges. We used an online survey to solicit feedback from wetland managers and biologists in the Southern Great Plains of North America to gain information on perceptions of wetland managers regarding climate change; assess how the effects of climate change are being addressed through management; and identify barriers to implementing climate change adaptation. The majority of wetland managers (63%) agreed they are currently experiencing effects of climate change in wetlands, and most respondents (76%) reported that changes in the timing of water availability throughout the year was the most likely impact. Managers reported using a diversity of approaches in managing for changing precipitation, with management of native and invasive plant species being the two most common practices. Lack of funding and personnel were the most commonly identified factors limiting manager's response to changing precipitation patterns. In addition, >50% of managers indicated uncertainty about the effects of climate change on wetlands as a barrier to management, which may relate to limited access to peer-reviewed science. While most of the management practices reported were short-term measures and may not reflect long-term adaptation for climate change, the fact that many managers are considering climate change in their management suggests that there is considerable opportunities to continue developing capacity for climate change adaptation in the region.

Why did the chicken not cross the road? Anthropogenic development influences the movement of a grassland bird.

Movement and selection are inherently linked behaviors that form the foundation of a species' space-use patterns. Anthropogenic development in natural ecosystems can result in a variety of behavioral responses that can involve changes in either movement (speed or direction of travel) or selection (resources used), which in turn may cause population-level consequences including loss of landscape connectivity. Understanding how a species alters these different behaviors in response to human activity is essential for effective conservation. In this study, we investigated the effects of anthropogenic development such as roads, power lines and oil wells on the greater prairie-chicken (Tympanuchus cupido) movement and selection behaviors in the post-nesting and non-breeding season. Our first objective was to assess using integrated step selection analysis (iSSA) if greater prairie-chickens altered their movement behaviors or their selection patterns when encountering oil wells, power lines, or roads. Our second objective was to determine whether prairie-chickens avoided crossing linear features such as roads or power lines by comparing the number of crossing events in greater prairie-chicken movement tracks to the number of movements that crossed these features in simulated movement tracks. Based on the iSSA analysis, we found that greater prairie-chickens avoided oil wells, power lines, and roads in both seasons, and altered their rate of movement when near anthropogenic structures. However, changes in speed varied by season, with prairie-chickens increasing their movement rates in the post-nesting season when near to development and decreasing movement rates in the non-breeding season. Furthermore, prairie-chickens crossed roads and power lines at much lower rates than expected. These changes in behavior can result in habitat loss for greater prairie-chickens, as well as the potential loss of landscape connectivity. By considering both movement and selection, we were able to develop an ecological understanding of how increasing human activity may influence the space use of this species of conservation concern. Furthermore, this research provides insight into the decision-making processes by animals when they encounter anthropogenic development.

Effects of pyric herbivory on prairie-chicken (Tympanuchus spp) habitat.

The reduction and simplification of grasslands has led to the decline of numerous species of grassland fauna, particularly grassland-obligate birds. Prairie-chickens (Tympanuchus spp.) are an example of obligate grassland birds that have declined throughout most of their distribution and are species of conservation concern. Pyric herbivory has been suggested as a land management strategy for enhancing prairie-chicken habitat and stabilizing declining population trends. We assessed differences in vegetation structure created by pyric herbivory compared to fire-only treatments to determine whether pyric herbivory increased habitat heterogeneity for prairie-chickens, spatially or temporally. Our study was performed at four sites in the southern Great Plains, all within the current or historic distribution of either lesser (T. pallidicinctus), greater (T. cupido), or Attwater's (T. cupido attwateri) prairie-chickens. Key vegetation characteristics of grass cover and vegetation height in pyric herbivory and fire-only treatments were within the recommended range of values for prairie-chickens during their distinct life history stages. However, patches managed via pyric herbivory provided approximately 5% more forb cover than fire-only treatments for almost 30 months post-fire. Additionally, pyric herbivory extended the length of time bare ground was present after fires. Pyric herbivory also reduced vegetation height and biomass, with mean vegetation height in pyric herbivory treatments lagging behind fire-only treatments by approximately 15 months. Canopy cover in fire-only treatments exceeded levels recommended for prairie-chicken young within 12 months post-fire. However, canopy cover in pyric herbivory treatments never exceeded the maximum recommended levels. Overall, it appears that pyric herbivory improves vegetation characteristics reported as critical to prairie-chicken reproduction. Based on our results, we suggest pyric herbivory as a viable management technique to promote prairie-chicken habitat in the southern Great Plains, while still accommodating livestock production.

Habitat associations of bats in a working rangeland landscape.

Land-use change has resulted in rangeland loss and degradation globally. These changes include conversion of native grasslands for row-crop agriculture as well as degradation of remaining rangeland due to fragmentation and changing disturbance regimes. Understanding how these and other factors influence wildlife use of rangelands is important for conservation and management of wildlife populations. We investigated bat habitat associations in a working rangeland in southeastern North Dakota. We used Petterson d500x acoustic detectors to systematically sample bat activity across the study area on a 1-km point grid. We identified calls using Sonobat autoclassification software. We detected five species using this working rangeland, which included Lasionycteris noctivagans (2,722 detections), Lasiurus cinereus (2,055 detections), Eptesicus fuscus (749 detections), Lasiurus borealis (62 detections), and Myotis lucifugus (1 detection). We developed generalized linear mixed-effects models for the four most frequently detected species based on their ecology. The activity of three bat species increased with higher tree cover. While the scale of selection varied between the four species, all three investigated scales were explanatory for at least one bat species. The broad importance of trees to bats in rangelands may put their conservation needs at odds with those of obligate grassland species. Focusing rangeland bat conservation on areas that were treed prior to European settlement, such as riparian forests, can provide important areas for bat conservation while minimizing negative impacts on grassland species.

Moderate patchiness optimizes heterogeneity, stability, and beta diversity in mesic grassland.

Heterogeneous disturbance patterns are fundamental to rangeland conservation and management because heterogeneity creates patchy vegetation, broadens niche availability, increases compositional dissimilarity, and enhances temporal stability of aboveground biomass production. Pyrodiversity is a popular concept for how variability in fire as an ecological disturbance can enhance heterogeneity, but mechanistic understanding of factors that drive heterogeneity is lacking. Mesic grasslands are examples of ecosystems in which pyrodiversity is linked strongly to broad ecological processes such as trophic interactions because grazers are attracted to recently burned areas, creating a unique ecological disturbance referred to as the fire-grazing interaction, or pyric herbivory. But several questions about the application of pyric herbivory remain: What proportion of a grazed landscape must burn, or how many patches are required, to create sufficient spatial heterogeneity and reduce temporal variability? How frequently should patches burn? Does season of fire matter? To bring theory into applied practice, we studied a gradient of grazed tallgrass prairie landscapes created by different sizes, seasons, and frequencies of fire, and used analyses sensitive to nonlinear trends. The greatest spatial heterogeneity and lowest temporal variability in aboveground plant biomass, and greatest plant functional group beta diversity, occurred in landscapes with three to four patches (25%-33% of area burned) and three- to four-year fire return intervals. Beta diversity had a positive association with spatial heterogeneity and negative relationship with temporal variability. Rather than prescribing that these results constitute best management practices, we emphasize the flexibility offered by interactions between patch number and fire frequency for matching rangeland productivity and offtake to specific management goals. As we observed no differences across season of fire, we recommend future research focus on fire frequency within a moderate proportion of the landscape burned, and consider a wider seasonal burn window.

Pyric-carnivory: Raptor use of prescribed fires.

Fire is a process that shaped and maintained most terrestrial ecosystems worldwide. Changes in land use and patterns of human settlement have altered fire regimes and led to fire suppression resulting in numerous undesirable consequences spanning individual species and entire ecosystems. Many obvious and direct consequences of fire suppression have been well studied, but several, albeit less obvious, costs of alteration to fire regimes on wildlife are unknown. One such phenomenon is the response of carnivores to fire events-something we refer to as pyric-carnivory. To investigate the prevalence of pyric-carnivory in raptors, we monitored 25 prescribed fires occurring during two different seasons and across two different locations in tallgrass prairie of the central United States. We used paired point counts occurring before and during prescribed fires to quantify the use of fires by raptors. We found a strong attraction to fires with average maximum abundance nearly seven times greater during fires than prior to ignitions (before: x¯ = 2.90, SE = 0.42; during: x¯ = 20.20; SE = 3.29) and an average difference between fire events and immediately before fires of 15.2 (±2.69) raptors. This result was driven by Swainson's hawks (Buteo swainsoni), which were the most abundant (n = 346) of the nine species we observed using fires. Our results illustrate the importance of fire as integral disturbance process that effects wildlife behavior through multiple mechanisms that are often overshadowed by the predominant view of fire as a tool used for vegetation management.

Effects of livestock grazing on rangeland biodiversity: A meta-analysis of grouse populations.

Livestock grazing affects over 60% of the world's agricultural lands and can influence rangeland ecosystem services and the quantity and quality of wildlife habitat, resulting in changes in biodiversity. Concomitantly, livestock grazing has the potential to be detrimental to some wildlife species while benefiting other rangeland organisms. Many imperiled grouse species require rangeland landscapes that exhibit diverse vegetation structure and composition to complete their life cycle. However, because of declining populations and reduced distributions, grouse are increasingly becoming a worldwide conservation concern. Grouse, as a suite of upland gamebirds, are often considered an umbrella species for other wildlife and thus used as indicators of rangeland health. With a projected increase in demand for livestock products, better information will be required to mitigate the anthropogenic effects of livestock grazing on rangeland biodiversity. To address this need, we completed a data-driven and systematic review of the peer-reviewed literature to determine the current knowledge of the effects of livestock grazing on grouse populations (i.e., chick production and population indices) worldwide. Our meta-analysis revealed an overall negative effect of livestock grazing on grouse populations. Perhaps more importantly, we identified an information void regarding the effects of livestock grazing on the majority of grouse species. Additionally, the reported indirect effects of livestock grazing on grouse species were inconclusive and more reflective of differences in the experimental design of the available studies. Future studies designed to evaluate the direct and indirect effects of livestock grazing on wildlife should document (i) livestock type, (ii) timing and frequency of grazing, (iii) duration, and (iv) stocking rate. Much of this information was lacking in the available published studies we reviewed, but is essential when making comparisons between different livestock grazing management practices and their potential impacts on rangeland biodiversity.

Reproductive plasticity and landscape heterogeneity benefit a ground-nesting bird in a fire-prone ecosystem.

Disturbance is critical for the conservation of rangeland ecosystems worldwide and many of these systems are fire dependent. Although it is well established that restoring fire as an ecological process can lead to increased biodiversity in grasslands and shrublands, the underlying mechanisms driving community patterns are poorly understood for fauna in fire-prone landscapes. Much of this uncertainty stems from the paucity of studies that examine the effects of fire at scales relevant to organism life histories. We assessed the response of a non-migratory ground-dwelling bird to disturbance (i.e., prescribed fire) and environmental stochasticity over the course of a 4-yr period, which spanned years of historic drought and record rainfall. Specifically, we investigated the nesting ecology of Northern Bobwhite (Colinus virginianus; hereafter Bobwhite) to illuminate possible avenues by which individuals respond to dynamic landscape patterns during a critical reproductive stage (i.e., nesting) in a mixed-grass shrubland in western Oklahoma, USA. We found that Bobwhites exhibited extreme plasticity in nest substrate use among time since fire categories (TSF) and subsequently maintained high nest survival (e.g., 57-70%). Bobwhites were opportunistic in nest substrate use among TSF categories (i.e., 72% of nest sites in shrubs in 0-12 months post fire compared to 71% in herbaceous vegetation in >36 months post fire), yet nesting decisions were first filtered by similar structural components (i.e., vertical and horizontal cover) within the vicinity of nest sites regardless of TSF category. Despite being a non-migratory and comparatively less mobile ground-nesting bird species, Bobwhites adjusted to dynamic vegetation mosaics on a fire-prone landscape under stochastic climatic conditions that culminated in stable and high nest survival. Broadly, our findings provide a unique depiction of organism response strategies to fire at scales relevant to a critical life-stage, a topic that has been previously understudied and poorly understood. We also demonstrate how doing so can better inform conservation practices aimed at restoring fire regimes on grassland and shrubland landscapes.

Restoring fire to grasslands is critical for migrating shorebird populations.

Fire is a disturbance process that maintains the structure and function of grassland ecosystems while sustaining grassland biodiversity. Conversion of grasslands to other land uses coupled with altered disturbance regimes has greatly diminished the habitat available to many grassland-dependent species. These changes have been linked to declines in breeding bird populations, but may also be critical for migrating bird populations such as those shorebird species that depend on mesic grasslands during migration. We examined migratory shorebird use of burned grasslands in the southern Great Plains of North America using DISTANCE sampling to estimate and compare bird densities across recently burned and not recently burned landscapes (1-5 yr post fire). We conducted two surveys per week for 8-10 weeks along a 54-km route starting at the end of March and concluding in mid-May during 2014-2015. We encountered 2,509 total shorebirds in recently burned areas compared to 130 individuals in areas that were unburned. Fire was a major attractant for our three focal species with American Golden-plover (Pluvialis dominica), Upland Sandpiper (Bartramia longicauda), and Killdeer (Charadrius vociferus) densities of 20.48, 11.09, and 26.09 birds/km2 in burned areas compared with 0.00, 1.27, and 0.92 birds/km2 in unburned areas, respectively. This research illustrates the importance of burned grassland for migrating shorebirds, a phenomenon that has largely gone unreported previously. Generally, these findings add to a body of knowledge that demonstrates the value of managing grasslands with historic disturbances that vary over space and time. The application of these findings should improve decision-making for shorebird conservation and provides evidence that prescribed fire planning should include consideration for breeding, transient, and non-breeding populations that vary in their temporal use of the landscape.

Temporal variability in aboveground plant biomass decreases as spatial variability increases.

Ecological theory predicts that diversity decreases variability in ecosystem function. We predict that, at the landscape scale, spatial variability created by a mosaic of contrasting patches that differ in time since disturbance will decrease temporal variability in aboveground plant biomass. Using data from a multi-year study of seven grazed tallgrass prairie landscapes, each experimentally managed for one to eight patches, we show that increased spatial variability driven by spatially patchy fire and herbivory reduces temporal variability in aboveground plant biomass. This pattern is associated with statistical evidence for the portfolio effect and a positive relationship between temporal variability and functional group synchrony as predicted by metacommunity variability theory. As disturbance from fire and grazing interact to create a shifting mosaic of spatially heterogeneous patches within a landscape, temporal variability in aboveground plant biomass can be dampened. These results suggest that spatially heterogeneous disturbance regimes contribute to a portfolio of ecosystem functions provided by biodiversity, including wildlife habitat, fuel, and forage. We discuss how spatial patterns of disturbance drive variability within and among patches.

Dynamic Disturbance Processes Create Dynamic Lek Site Selection in a Prairie Grouse.

It is well understood that landscape processes can affect habitat selection patterns, movements, and species persistence. These selection patterns may be altered or even eliminated as a result of changes in disturbance regimes and a concomitant management focus on uniform, moderate disturbance across landscapes. To assess how restored landscape heterogeneity influences habitat selection patterns, we examined 21 years (1991, 1993-2012) of Greater Prairie-Chicken (Tympanuchus cupido) lek location data in tallgrass prairie with restored fire and grazing processes. Our study took place at The Nature Conservancy's Tallgrass Prairie Preserve located at the southern extent of Flint Hills in northeastern Oklahoma. We specifically addressed stability of lek locations in the context of the fire-grazing interaction, and the environmental factors influencing lek locations. We found that lek locations were dynamic in a landscape with interacting fire and grazing. While previous conservation efforts have treated leks as stable with high site fidelity in static landscapes, a majority of lek locations in our study (i.e., 65%) moved by nearly one kilometer on an annual basis in this dynamic setting. Lek sites were in elevated areas with low tree cover and low road density. Additionally, lek site selection was influenced by an interaction of fire and patch edge, indicating that in recently burned patches, leks were located near patch edges. These results suggest that dynamic and interactive processes such as fire and grazing that restore heterogeneity to grasslands do influence habitat selection patterns in prairie grouse, a phenomenon that is likely to apply throughout the Greater Prairie-Chicken's distribution when dynamic processes are restored. As conservation moves toward restoring dynamic historic disturbance patterns, it will be important that siting and planning of anthropogenic structures (e.g., wind energy, oil and gas) and management plans not view lek locations as static points, but rather as sites that shift around the landscape in response to shifting vegetation structure. Acknowledging shifting lek locations in these landscapes will help ensure conservation efforts are successful by targeting the appropriate areas for protection and management.

Predicting Greater Prairie-Chicken Lek Site Suitability to Inform Conservation Actions.

The demands of a growing human population dictates that expansion of energy infrastructure, roads, and other development frequently takes place in native rangelands. Particularly, transmission lines and roads commonly divide rural landscapes and increase fragmentation. This has direct and indirect consequences on native wildlife that can be mitigated through thoughtful planning and proactive approaches to identifying areas of high conservation priority. We used nine years (2003-2011) of Greater Prairie-Chicken (Tympanuchus cupido) lek locations totaling 870 unique leks sites in Kansas and seven geographic information system (GIS) layers describing land cover, topography, and anthropogenic structures to model habitat suitability across the state. The models obtained had low omission rates (<0.18) and high area under the curve scores (AUC >0.81), indicating high model performance and reliability of predicted habitat suitability for Greater Prairie-Chickens. We found that elevation was the most influential in predicting lek locations, contributing three times more predictive power than any other variable. However, models were improved by the addition of land cover and anthropogenic features (transmission lines, roads, and oil and gas structures). Overall, our analysis provides a hierarchal understanding of Greater Prairie-Chicken habitat suitability that is broadly based on geomorphological features followed by land cover suitability. We found that when land features and vegetation cover are suitable for Greater Prairie-Chickens, fragmentation by anthropogenic sources such as roadways and transmission lines are a concern. Therefore, it is our recommendation that future human development in Kansas avoid areas that our models identified as highly suitable for Greater Prairie-Chickens and focus development on land cover types that are of lower conservation concern.

Spatial heterogeneity increases diversity and stability in grassland bird communities.

Grasslands are inherently dynamic in space and time, evolving with frequent disturbance from fire and herbivores. As a consequence of human actions, many remaining grasslands have become homogenous, which has led to reduced ecosystem function, biodiversity loss, and decreased ecological services. Previous research has shown that restoring inherent heterogeneity to grasslands can increase avian diversity, but the amount of heterogeneity (i.e., number of patches or fire return interval) and the impact on avian community stability have yet to be investigated. We used a unique landscape-level design to examine avian response to interacting fire and grazing across multiple experimental landscapes that represented a gradient of fire- and grazing-dependent heterogeneity. We used seven landscapes (430-980 ha; x = 627 ha) with varying levels of patchiness ranging from annually burned (one single patch) with spring-only fires to a four-year fire return interval with spring and summer fires (eight patches). This design created a range of heterogeneity as a result of pyric herbivory, an ecological process in which fire and grazing are allowed to interact in space and time. We found that greater heterogeneity across experimental landscapes resulted in increased avian diversity and stability over time. An index of bird community change, quantified as the sum of the range of detrended correspondence analysis axis site scores, was nearly four times greater in the most homogenous experimental landscape when compared to the most heterogeneous experimental landscape. Species responses were consistently positively associated with increased heterogeneity at the landscape scale, and within-experimental-landscape responses were most often related to litter cover, litter accumulation, and vegetation height. We conclude that increased fire- and grazig-dependent heterogeneity can result in high variability in the bird community at finer, transect scales, but increased diversity and stability at broad landscape scales. We recommend that future management efforts in rangelands focus on restored disturbance processes to increase heterogeneity and improve grassland bird conservation.

Conservation implications of native and introduced ungulates in a changing climate.

In many grasslands, grazing by large native or introduced ungulates drives ecosystem structure and function. The behavior of these animals is important as it directs the spatial effects of grazing. To the degree that temperature drives spatial components of foraging, understanding how changes in climate alter grazing behavior will provide guidance for the conservation of ecosystem goods and services. We determined the behavioral response of native bison (Bison bison) and introduced cattle (Bos taurus) to temperature in tallgrass prairie within the Great Plains, USA. We described the thermal environment by measuring operative temperature (the temperature perceived by animals) through space and time. Site selection preferences of ungulates were quantified using resource selection functions. Woody vegetation in tallgrass prairie provided a cooler thermal environment for large ungulates, decreasing operative temperature up to 16 °C in the heat of the summer. Cattle began to seek thermal refugia at lower air temperatures (24 °C) by selecting areas closer to woody vegetation and water sources. Bison, however, sought refugia within wooded areas at higher air temperatures (36 °C), which occurred much less frequently. Both species became more attracted to riparian areas as air temperature increased, with preferences increasing tenfold during the hottest periods. As predicted warming occurs across the Great Plains and other grasslands, grazing behavior and subsequent grazing effects will be altered. Riparian areas, particularly those with both water and woody vegetation, will receive greater utilization and selection by large ungulates. The use of native grazers for conservation or livestock production may mitigate negative effects caused by increased temperatures.