Factors influencing the persistence of a fire-sensitive Artemisia species in a fire-dependent ecosystem.

Fire refugia and patchiness are important to the persistence of fire-sensitive species and may facilitate biodiversity conservation in fire-dependent landscapes. Playing the role of ecosystem engineers, large herbivores alter vegetation structure and can reduce wildfire risk. However, herbivore effects on the spatial variability of fire and the persistence of fire-sensitive species are not clear. To examine the hypothesis that large herbivores support the persistence of fire-sensitive species through the creation of fire refugia in fire-prone landscapes, we examined the response of a fire-sensitive plant, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis [Beetle & Young]) to fire and grazing in the fire-dependent mixed-grass prairie of the northern Great Plains. We carried out a controlled burn in 2010 within pre-established exclosures that allowed differential access to wild and domestic herbivores and no record of fire in the previous 75 years due to fire suppression efforts. The experiment was set up with a split-plot design to also examine potential changes in plots that were not burned. Canopy cover of big sagebrush was recorded before the burn in 2010 and again in 2011 with percent area burned recorded within 1-month post-fire in the burned plots. Percentage area burned was the greatest in ungulate exclosures (92% ± 2%) and the least in open areas (55% ± 21%), suggesting that large herbivores influenced fire behavior (e.g., reducing fire intensity and rate of spread) and are likely to increase fire patchiness through their alterations to the fuel bed. Regression analysis indicated that the proportion of sagebrush cover lost was significantly correlated with the proportion of area burned (R2  = 0.76, p = 0.05). No differences in the non-burn plots were observed among grazing treatments or among years. Altogether, this illustrates the potential importance of large herbivores in creating biotic-driven fire refugia for fire-sensitive species to survive within the flammable fuel matrix of fire-dependent grassland ecosystems such as the mixed-grass prairie. Our findings also attest to the resiliency of the northern Great Plains to fire and herbivory and underscore the value of managing grasslands for heterogeneity with spatial and temporal variations in these historic disturbances.

Fire management alters the thermal landscape and provides multi-scale thermal options for a terrestrial turtle facing a changing climate.

As effects of climate change intensify, there is a growing need to understand the thermal properties of landscapes and their influence on wildlife. A key thermal property of landscapes is vegetation structure and composition. Management approaches can alter vegetation and consequently the thermal landscape, potentially resulting in underappreciated consequences for wildlife thermoregulation. Consideration of spatial scale can clarify how management overlaid onto existing vegetation patterns affects thermal properties of landscapes relevant to wildlife. We examined effects of temperature, fire management, and vegetation structure on multi-scale habitat selection of an ectothermic vertebrate (the turtle Terrapene carolina triunguis) in the Great Plains of the central United States by linking time-since-fire data from 18 experimental burn plots to turtle telemetry locations and thermal and vegetation height data. Within three 60-ha experimental landscapes, each containing six 10-ha sub-blocks that are periodically burned, we found that turtles select time-since-fire gradients differently depending on maximum daily ambient temperature. At moderate temperatures, turtles selected sub-blocks with recent (<1 year) time-since-fire, but during relatively hot and cool conditions, they selected sub-blocks with later (2-3 year) time-since-fire that provided thermal buffering compared with recently burned sub-blocks. Within 10-ha sub-blocks, turtles selected locations with taller vegetation during warmer conditions that provided thermal buffering. Thermal performance curves revealed that turtle activity declined as temperatures exceeded ~24-29°C, and on "heat days" (≥29°C) 73% of turtles were inactive compared with 37% on non-heat days, emphasizing that thermal extremes may lead to opportunity costs (i.e., foregone benefits turtles could otherwise accrue if active). Our results indicate that management approaches that promote a mosaic of vegetation heights, like spatiotemporally dynamic fire, can provide thermal refuges at multiple spatial scales and thus be an actionable way to provide wildlife with multiple thermal options in the context of ongoing and future climate change.

Role of the thermal environment in scaled quail (Callipepla squamata) nest site selection and survival.

Temperature is increasingly recognized as an important component of wildlife habitat. Temperature is particularly important for avian nest sites, where extreme temperatures can influence adult behavior, embryonic development, and survival. For species inhabiting arid and semiarid climates, such as the scaled quail (Callipepla squamata), frequent exposure to extreme temperatures may increase the importance of the nest microclimate. Limited data suggest that scaled quail respond to temperature when selecting nest sites, and they are also known to respond to the presence of surface water and shrub cover on the landscape, two resources which may mitigate thermal stress. To better understand the role of temperature in nest site selection and survival, and to evaluate how other landscape resources may benefit nesting quail, we investigated nest site characteristics of scaled quail in southeastern New Mexico, USA. During the breeding seasons of 2018 and 2019 we located nests, monitored nest fate, and recorded thermal and vegetation characteristics at three spatial scales: the nest bowl, the nest microsite (area within 10 m of the nest bowl), and the landscape. We found that nest bowls moderated temperature relative to both the surrounding microsite and the broader landscape, remaining almost 5 °C cooler on average than the surrounding microsite at mid-day. Nest bowls also had taller, greater cover of vegetation compared to both the surrounding microsites and the landscape. Despite apparent selection for cooler temperatures and taller vegetation, these characteristics demonstrated a weak relationship with nest survival. Rather, nest survival was positively influenced by proximity to surface water and honey mesquite (Prosopis glandulosa), with survival decreasing with increasing distance from these features. Although the mechanism for this relationship is unclear, our results support the importance of temperature for nest site selection of ground-nesting birds in semiarid landscapes, and suggest further exploration of landscape-level sources of thermal mitigation.

The effects of fire on the thermal environment of sagebrush communities.

Thermal heterogeneity provides options for organisms during extreme temperatures that can contribute to their fitness. Sagebrush (Artemisia spp.) communities exhibit vegetation heterogeneity that creates thermal variation at fine spatial scales. However, fire can change vegetation and thereby variation within the thermal environment of sagebrush communities. To describe spatial and temporal thermal variation of sagebrush communities following wildfire, we measured black bulb temperature (Tbb) at 144 random points dispersed within unburned and burned communities, for 24-h at each random point. We observed a wide thermal gradient in unburned (-7.3° to 63.3 °C) and burned (-4.6° to 64.8 °C) sagebrush communities. Moreover, unburned and burned sagebrush communities displayed high thermal heterogeneity relative to ambient temperature (Tair). Notably, Tbb varied by 47 °C in both unburned and burned communities when Tair was 20 °C. However, fire greatly reduced the buffering capacity and thermal refuge of Wyoming big sagebrush (A. tridentata wyomingensis) communities during low and high Tair. Furthermore, fire increased Tbb in Wyoming big sagebrush and mountain big sagebrush (A. t. vaseyana) during the mid-day hours. These results demonstrate how fire changes the thermal environment of big sagebrush communities and the importance of shrub structure which can provide thermal refuge for organisms in burned communities during extreme low and high Tair.

Behavioral modifications lead to disparate demographic consequences in two sympatric species.

Life-history theory suggests species that typically have a large number of offspring and high adult mortality may make decisions that benefit offspring survival in exchange for increased adult risks. Such behavioral adaptations are essential to understanding how demographic performance is linked to habitat selection during this important life-history stage. Though studies have illustrated negative fitness consequences to attendant adults or potential fitness benefits to associated offspring because of adaptive habitat selection during brood rearing, equivocal relationships could arise if both aspects of this reproductive trade-off are not assessed simultaneously. To better understand how adaptive habitat selection during brood rearing influences demographics, we studied the brood survival, attendant parental survival, and space use of two sympatric ground-nesting bird species, the northern bobwhite (hereafter: "bobwhite"; Colinus virgininanus) and scaled quail (Callipepla squamata). During the 2013-2014 breeding seasons, we estimated habitat suitability across two grains (2 m and 30 m) for both species and determined how adult space use of these areas influenced individual chick survival and parental risk. We found the proportion of a brood's home range containing highly suitable areas significantly increased bobwhite chick survival (ß = 0.02, SE = 0.006). Additionally, adult weekly survival for bobwhite was greater for individuals not actively brooding offspring (0.9716, SE = 0.0054) as compared to brooding adults (0.8928, SE = 0.0006). Conversely, brood habitat suitability did not influence scaled quail chick survival during our study, nor did we detect a survival cost for adults that were actively brooding offspring. Our research illustrates the importance of understanding life-history strategies and how they might influence relationships between adaptive habitat selection and demographic parameters.

Thermal refuge affects space use and movement of a large-bodied galliform.

Temperature affects every organism on Earth and has been argued to be one of the most critical factors influencing organisms' ecology and evolution. Most organisms are susceptible to landscape temperature ranges that exceed their thermal tolerance. As a result, the distribution of landscape features that mitigate thermal extremes can affect movement and space use of organisms. Using Rio Grande wild turkey (Meleagris gallopavo intermedia) as a model species, we measured black bulb temperature throughout the diurnal period and identified vegetation characteristics at wild turkey locations and random landscape locations. We observed that the thermal landscape was highly heterogeneous with temperature varying up to 52 °C at a given ambient temperature. Vegetation type strongly influenced temperature across space during daily peak heating, with taller vegetation types (woody vegetation >2 m) having mean temperatures up to 8.95 °C cooler than the remainder of the landscape. However, these cooler vegetation types were uncommon, only accounting for 8.2% of the landscape. Despite the rarity of tall woody cover, wild turkey showed strong selection for this vegetation type particularly during peak daily heating with 74.9% of locations within 18 m of tree cover. Not only did wild turkey alter space use across time relative to temperature variation, but they also altered movement. We found that on the hottest days (≥35 °C), wild turkeys decreased movement by three fold during peak heating, while movement on cooler days (<30 °C) was uniform. Collectively, our data provide evidence that space use and movement for large avian species can be influenced by the thermal environment, and that the thermal environment is an important component of habitat for a species.

Perspectives on grassland conservation efforts: should we rewild to the past or conserve for the future?

Grasslands are among the most imperilled biomes of the world. Identifying the most appropriate framework for restoring grasslands is dependent on the objectives of restoration, which is inherently determined by human priorities. Debates over the appropriate conservation model for grasslands have often focused on which species of herbivores should be the focus of restoration efforts. Here we discuss three perspectives of herbivore-based conservation in North American grasslands. First, the Pleistocene rewilding perspective is based upon the idea that early humans contributed to the demise of megafauna that were important to the evolution and development of many of North America's grasslands; therefore, their aim of restoration is rewilding of landscapes to pre-human times. Second, the bison rewilding perspective considers American bison a keystone herbivore that is culturally and ecologically important to North American grasslands. A third perspective focuses on restoring the pattern and processes of herbivory on grasslands and is less concerned about which herbivore is introduced to the landscape. We evaluate each of these three conservation perspectives in terms of a framework that includes a human domain, an herbivore domain and a biophysical domain. While all conservation perspectives partly address the three domains, they all fall short in key areas. Specifically, they fail to recognize that past, current and future humans are intimately linked to grassland patterns and processes and will continue to play a role in structuring grasslands. Furthermore, these perspectives seem to only superficially consider the role of fragmentation and climate change in influencing grassland patterns and processes. As such, we argue that future grassland conservation efforts must depend on the development of a model that better integrates societal, economic and policy objectives and recognizes climate change, fragmentation and humans as an integral part of these ecosystems.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Avian parental behavior and nest success influenced by temperature fluctuations.

Behavioral adjustments and parental decisions during reproduction can influence the thermal environment at nests, yet our understanding into how environmental factors (i.e., temperature and precipitation) constrain an adult's ability to balance self-maintenance and incubation demands is limited. To expand our understanding of how species respond to environmental factors, we investigated the reproductive ecology of two ground-nesting species (northern bobwhite [Colinus virginianus] and scaled quail [Callipepla squamata]) in a region (i.e., the Southern Great Plains) prone to thermal variability (i.e., extreme hot and cold temperatures). Specifically, our objective was to examine how temperature and precipitation directly influenced behavioral adjustments (i.e., off-bout duration, frequency, and nest attentiveness) and parental decisions (i.e., nest site selection), and indirectly influenced nest fate. Overall, we found that parents chose to nest in sites that were significantly cooler in temperature than randomly selected sites, and parents further altered the thermal environment experienced by embryos through incubation behavior. Daily precipitation and average ambient temperature and/or their interaction best predicted incubation behaviors, yet each species differed in the timing (i.e., morning vs. evening), frequency, and duration of off-bouts. Furthermore, successful nests were associated with cooler nest site temperatures for bobwhite and warmer nest site temperatures for scaled quail. Our finding of relatively stable (35.5 °C) incubation temperature for developing embryos of both species suggests that ground-nesting birds are able to regulate microclimate through behavioral adjustments and parental decisions even under extreme temperature fluctuations. Nevertheless, the ability for a ground-nesting species to effectively modify behavioral adjustments and decisions may be altered during long periods of enhanced physiological and environmental stress.

Variation and drivers of airflow patterns associated with olfactory concealment and habitat selection.

Many terrestrial predators rely on olfaction to detect prey; therefore, prey should select habitat to reduce detectability of their odor cues. One way prey can potentially conceal their odor is by selecting locations with high turbulence and/or updrafts, conditions that disperse odor plumes and make odor sources difficult to locate. However, it is unclear how these conditions vary among vegetation cover types and which vegetative features drive them. We assessed variation and drivers of variables hypothesized to influence olfactory concealment (turbulence intensity and airflow slope) and experimentally evaluated whether these variables indeed influence predator detection of simulated prey. Specifically, we compared vegetation patch-scale values of turbulence intensity and airflow slope among grassland, shrubland, and forest and assessed relationships among these airflow variables and local-scale vegetative features within each vegetation type. Additionally, we experimentally investigated the importance of turbulence intensity, airflow slope, and visual concealment for predicting predator detection of scented quail eggs. In all vegetation types, we documented high variability in airflow conditions. At the patch scale, turbulence intensity was greater in shrubland and grassland than in forest, and updrafts were most common in shrubland whereas downdrafts were most common in grassland. Grassland was the only vegetation type with strong relationships among turbulence intensity and local vegetation features; both visual concealment and vegetation height were positively related to turbulence intensity. Additionally, persistence of scented quail eggs in grassland was best predicted by turbulence intensity; egg persistence increased with turbulence intensity. Our characterization of differences in olfactory variables among vegetation types provides an important step towards building a clearer understanding of olfactory landscapes. Further, our observation of both patch- and local-scale variation in olfactory variables suggests that prey can potentially select for olfactory concealment at multiple scales. We hypothesize that olfactory concealment provided by high levels of turbulence intensity is an important component of cover in grassland, and that in grassland/shrubland landscapes, prey selection of shrubland patches reduces odor detectability. Our finding of a positive relationship between turbulence intensity and visual concealment also suggests that olfactory concealment may be a previously underappreciated confounding factor in studies of habitat selection.

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.

Incorporating abundance information and guiding variable selection for climate-based ensemble forecasting of species' distributional shifts.

Ecological niche models (ENMs) have increasingly been used to estimate the potential effects of climate change on species' distributions worldwide. Recently, predictions of species abundance have also been obtained with such models, though knowledge about the climatic variables affecting species abundance is often lacking. To address this, we used a well-studied guild (temperate North American quail) and the Maxent modeling algorithm to compare model performance of three variable selection approaches: correlation/variable contribution (CVC), biological (i.e., variables known to affect species abundance), and random. We then applied the best approach to forecast potential distributions, under future climatic conditions, and analyze future potential distributions in light of available abundance data and presence-only occurrence data. To estimate species' distributional shifts we generated ensemble forecasts using four global circulation models, four representative concentration pathways, and two time periods (2050 and 2070). Furthermore, we present distributional shifts where 75%, 90%, and 100% of our ensemble models agreed. The CVC variable selection approach outperformed our biological approach for four of the six species. Model projections indicated species-specific effects of climate change on future distributions of temperate North American quail. The Gambel's quail (Callipepla gambelii) was the only species predicted to gain area in climatic suitability across all three scenarios of ensemble model agreement. Conversely, the scaled quail (Callipepla squamata) was the only species predicted to lose area in climatic suitability across all three scenarios of ensemble model agreement. Our models projected future loss of areas for the northern bobwhite (Colinus virginianus) and scaled quail in portions of their distributions which are currently areas of high abundance. Climatic variables that influence local abundance may not always scale up to influence species' distributions. Special attention should be given to selecting variables for ENMs, and tests of model performance should be used to validate the choice of variables.

Influence of olfactory and visual cover on nest site selection and nest success for grassland-nesting birds.

Habitat selection by animals is influenced by and mitigates the effects of predation and environmental extremes. For birds, nest site selection is crucial to offspring production because nests are exposed to extreme weather and predation pressure. Predators that forage using olfaction often dominate nest predator communities; therefore, factors that influence olfactory detection (e.g., airflow and weather variables, including turbulence and moisture) should influence nest site selection and survival. However, few studies have assessed the importance of olfactory cover for habitat selection and survival. We assessed whether ground-nesting birds select nest sites based on visual and/or olfactory cover. Additionally, we assessed the importance of visual cover and airflow and weather variables associated with olfactory cover in influencing nest survival. In managed grasslands in Oklahoma, USA, we monitored nests of Northern Bobwhite (Colinus virginianus), Eastern Meadowlark (Sturnella magna), and Grasshopper Sparrow (Ammodramus savannarum) during 2015 and 2016. To assess nest site selection, we compared cover variables between nests and random points. To assess factors influencing nest survival, we used visual cover and olfactory-related measurements (i.e., airflow and weather variables) to model daily nest survival. For nest site selection, nest sites had greater overhead visual cover than random points, but no other significant differences were found. Weather variables hypothesized to influence olfactory detection, specifically precipitation and relative humidity, were the best predictors of and were positively related to daily nest survival. Selection for overhead cover likely contributed to mitigation of thermal extremes and possibly reduced detectability of nests. For daily nest survival, we hypothesize that major nest predators focused on prey other than the monitored species' nests during high moisture conditions, thus increasing nest survival on these days. Our study highlights how mechanistic approaches to studying cover informs which dimensions are perceived and selected by animals and which dimensions confer fitness-related benefits.

Using a historic drought and high-heat event to validate thermal exposure predictions for ground-dwelling birds.

Deviations from typical environmental conditions can provide insight into how organisms may respond to future weather extremes predicted by climate modeling. During an episodic and multimonth heat wave event (i.e., ambient temperature up to 43.4°C), we studied the thermal ecology of a ground-dwelling bird species in Western Oklahoma, USA. Specifically, we measured black bulb temperature (Tbb) and vegetation parameters at northern bobwhite (Colinus virginianus; hereafter bobwhite) adult and brood locations as well as at stratified random points in the study area. On the hottest days (i.e., ≥39°C), adults and broods obtained thermal refuge using tall woody cover that remained on average up to 16.51°C cooler than random sites on the landscape which reached >57°C. We also found that refuge sites used by bobwhites moderated thermal conditions by more than twofold compared to stratified random sites on the landscape but that Tbb commonly exceeded thermal stress thresholds for bobwhites (39°C) for several hours of the day within thermal refuges. The serendipitous high heat conditions captured in our study represent extreme heat for our study region as well as thermal stress for our study species, and subsequently allowed us to assess ground-dwelling bird responses to temperatures that are predicted to become more common in the future. Our findings confirm the critical importance of tall woody cover for moderating temperatures and functioning as important islands of thermal refuge for ground-dwelling birds, especially during extreme heat. However, the potential for extreme heat loads within thermal refuges that we observed (albeit much less extreme than the landscape) indicates that the functionality of tall woody cover to mitigate heat extremes may be increasingly limited in the future, thereby reinforcing predictions that climate change represents a clear and present danger for these species.

Factors Affecting Public Preferences for Grassland Landscape Heterogeneity in the Great Plains.

Agricultural intensification has fragmented rangelands in the Great Plains, which has contributed to uniform and homogeneous landscapes and decreased biodiversity. Alternative land management practices involving fire-grazing interactions can help maintain biodiversity without affecting livestock productivity. A survey was designed to understand the factors that influence preferences among the general population towards grassland landscape heterogeneity. Given the ordinal nature of survey responses, requisite data were analyzed using a generalized ordinal logit model. Results suggested that respondents who valued open space and those who recognized a need for a varying mix of uniform grasses and grasslands preferred landscape heterogeneity. Female respondents were about two times as likely to prefer heterogeneous landscapes compared to male respondents. In contrast, population groups that preferred wildlife habitat did not desire heterogeneous landscapes. Results suggest the need for extension and outreach activities to educate certain segments of the general population regarding benefits of alternative management practices that support landscape heterogeneity in the Great Plains.

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.

Extreme climatic events constrain space use and survival of a ground-nesting bird.

Two fundamental issues in ecology are understanding what influences the distribution and abundance of organisms through space and time. While it is well established that broad-scale patterns of abiotic and biotic conditions affect organisms' distributions and population fluctuations, discrete events may be important drivers of space use, survival, and persistence. These discrete extreme climatic events can constrain populations and space use at fine scales beyond that which is typically measured in ecological studies. Recently, a growing body of literature has identified thermal stress as a potential mechanism in determining space use and survival. We sought to determine how ambient temperature at fine temporal scales affected survival and space use for a ground-nesting quail species (Colinus virginianus; northern bobwhite). We modeled space use across an ambient temperature gradient (ranging from -20 to 38 °C) through a maxent algorithm. We also used Andersen-Gill proportional hazard models to assess the influence of ambient temperature-related variables on survival through time. Estimated available useable space ranged from 18.6% to 57.1% of the landscape depending on ambient temperature. The lowest and highest ambient temperature categories (<-15 °C and >35 °C, respectively) were associated with the least amount of estimated useable space (18.6% and 24.6%, respectively). Range overlap analysis indicated dissimilarity in areas where Colinus virginianus were restricted during times of thermal extremes (range overlap = 0.38). This suggests that habitat under a given condition is not necessarily a habitat under alternative conditions. Further, we found survival was most influenced by weekly minimum ambient temperatures. Our results demonstrate that ecological constraints can occur along a thermal gradient and that understanding the effects of these discrete events and how they change over time may be more important to conservation of organisms than are average and broad-scale conditions as typically measured in ecological studies.

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.

A Ground-Nesting Galliform's Response to Thermal Heterogeneity: Implications for Ground-Dwelling Birds.

The habitat selection choices that individuals make in response to thermal environments influence both survival and reproduction. Importantly, the way that organisms behaviorally respond to thermal environments depends on the availability and juxtaposition of sites affording tolerable or preferred microclimates. Although, ground nesting birds are especially susceptible to heat extremes across many reproductive stages (i.e., breeding, nesting, brood rearing), the mechanistic drivers of nest site selection for these species are not well established from a thermal perspective. Our goal was to assess nest site selection relative to the configuration of the thermal landscape by quantifying thermal environments available to a ground-nesting bird species inhabiting a climatically stressful environment. Using northern bobwhite (Colinus virginanus) as a model species, we measured black bulb temperature (Tbb) and vegetation parameters at 87 nests, 87 paired sites and 205 random landscape sites in Western Oklahoma during spring and summer 2013 and 2014. We found that thermal space within the study area exhibited differences in Tbb of up to 40°C during peak diurnal heating, resulting in a diverse thermal landscape available to ground-nesting birds. Within this thermally heterogeneous landscape, nest sites moderated Tbb by more than 12°C compared to random landscape sites. Furthermore, successful nests remained on average 6°C cooler than unsuccessful nests on days experiencing ambient temperatures ≥ 39°C. Models of future Tbb associated with 2080 climate change projections indicate that nesting bobwhites will face substantially greater Tbb throughout the landscape for longer durations, placing an even greater importance on thermal choices for nest sites in the future. These results highlight the capacity of landscape features to act as moderators of thermal extremes and demonstrate how thermal complexity at organism-specific scales can dictate habitat selection.

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.