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Toxicological effects assessment for wildlife in the 21st Century: Review of current methods and recommendations for a path forward

Model species (e.g., granivorous gamebirds, waterfowl, passerines, domesticated rodents) have been used for decades in guideline laboratory tests to generate survival, growth and reproductive data for prospective Ecological Risk Assessments (ERAs) for birds and mammals, while officially adopted risk assessment schemes for amphibians and reptiles do not exist. There are recognized shortcomings of current in vivo methods as well as uncertainty around the extent to which species with different life histories (e.g., terrestrial amphibians, reptiles, bats) than these commonly used models are protected by existing ERA frameworks. Approaches other than validating additional animal models for testing are being developed, but incorporation of such new approach methodologies (NAMs) into risk assessment frameworks will require robust validations against in vivo responses. This takes time, and the ability to extrapolate findings from non-animal studies to organism- and population-level effects in terrestrial wildlife remains weak. Failure to adequately anticipate and predict hazards could have economic and potentially even legal consequences for regulators and product registrants. In order to be able to use fewer animals or replace them altogether in the long-term, vertebrate use and whole organism data will be needed to provide data for NAMs validation in the short term. Therefore, it is worth investing resources for potential updates to existing standard test guidelines used in the laboratory as well as addressing the need for clear guidance on conduct of field studies. Herein we review the potential for improving standard in vivo test methods and for advancing the use of field studies in wildlife risk assessment, as these tools will be needed into the foreseeable future.

Integrated Environmental Assessment and Management↗

Survival of female mallards along the Vermont-Quebec border region

Understanding effects of location and timing of harvest seasons on mortality of ducks and geese from hunting is important in forming regulations that sustain viable waterfowl populations throughout their range. During 1990 and 1991 we alternately marked 80 hatching year (HY), female mallards along the Vermont–Quebec border; half with radio-transmitters and bands and half with only aluminum leg bands. We monitored radio-marked ducks daily and recorded survival status weekly for 15 weeks from August until December each year. Mallard mortalities began 25 September when the hunting season opened in the Province of Quebec, Canada. Overall survival of mallards at week 10 did not differ between years (0.51 in 1990 vs. 0.43 in 1991) or differ from that of HY American black ducks (0.44 females, 0.42 males) based on proportional hazard analysis in a Bayesian framework. The mortality rates for mallards from hunting (0.47) and causes unrelated to hunting (0.06) were similar between years and to those rates for HY black ducks at that same site. Hunter harvest accounted for most of the mortality recorded during this study and illegal feeding (i.e., baiting) at sites just before and during the hunting season was observed. Females with lower body condition index had greater mortality rates; a 1-standard-deviation increase in condition index would reduce mortality hazard by about 29%. Management options that may increase mallard survival in the area include lowering daily bag limit in Quebec and suspending split hunting seasons in Vermont that increase harvest, delaying opening date of hunting in Quebec to allow for increased body condition before hunting season opens, and improving enforcement of baiting restrictions.

Quebec, Vermont↗

Temporal variation in survival and recovery rates of lesser scaup: A response

We recently analyzed long-term (1951–2011) continental band-recovery data from lesser scaup ( Aythya affinis ) and demonstrated that harvest rates declined through time, but annual survival rates exhibited no such trends; moreover, annual harvest and survival rates were uncorrelated for all age-sex classes. We therefore concluded that declining fecundity was most likely responsible for recent population declines, rather than changes in harvest or survival. Lindberg et al. ( 2017 ) critiqued our conclusions, arguing that we did little more than fail to reject a null hypothesis of compensatory mortality, postulated ecologically unrealistic changes in fecundity, and failed to give sufficient consideration to additive harvest mortality. Herein, we re-summarize our original evidence indicating that harvest has been compensatory, or at most weakly additive, and demonstrate that our analysis had sufficient power to detect strongly additive mortality if it occurred. We further demonstrate that our conclusions were not confounded by population size, band loss, or individual heterogeneity, as suggested by Lindberg et al. ( 2017 ), and we provide additional support for our conjecture that low fecundity played a major role in declining scaup populations during 1983–2006. We therefore reiterate our original management recommendations: given low harvest rates and lack of demonstrable effect on scaup survival, harvest regulations could return to more liberal frameworks, and waterfowl biologists should work together to continue banding lesser scaup and use these data to explore alternative hypotheses to identify the true ecological causes of population change, given that it is unlikely to be excessive harvest.

Journal of Wildlife Management↗

Duck nest depredation, predator behavior, and female response using video

Depredation plays an important role in determining duck nest success and predator and female duck behavior during nest depredation can influence nest fate. We examined depredation of mallard ( Anas platyrhynchos ) and gadwall ( A. strepera ) nests in Suisun Marsh, California, USA, in 2015–2016 with continuous infrared video monitoring to identify nest predators and characterize predator and female duck behavior during depredation events. We recorded predators at 44% of 147 nests monitored. Raccoons ( Procyon lotor ) were the most frequent predator observed at nests (40% of nests visited and 53% of depredated eggs) followed by striped skunks ( Mephitis mephitis ; 27% and 27%), coyotes ( Canis latrans ; 4% and 9%), common ravens ( Corvus corax ; 4% and 9%), gopher snakes ( Pituophis catenifer catenifer ; 19% and 0%), and western yellow‐bellied racers ( Coluber constrictor mormon ; 1% and 0%). The number of eggs depredated per depredation bout varied among predators (raccoons: 7.3 eggs; skunks: 2.5; coyotes: 7.4; ravens: 7.7; and snakes: 0.0). Mammal depredation occurred between 1600 and 0400, whereas snakes and ravens were observed at nests during the day (snakes: 1000–2100; ravens: 1700). Females flushed from nests immediately before predator arrival ( = 29.0 ± 16.6 [SD] sec), and this timing did not vary among predators. However, the length of nest depredation bouts varied among predators. Nest visits by gopher snakes were longer (18.6 ± 19.4 min) than depredation bouts by other predators (12.3 ± 10.7 min), but snakes did not successfully consume any eggs. Females took more time to return to nests when nests were depredated by raccoons (239 ± 137 min), whereas females returned more quickly when nests were visited by skunks (81 ± 163 min) or gopher snakes (15 ± 128 min). Partial clutch depredation occurred at 15% of depredated nests, but only 23% of partially depredated nests successfully hatched ≥1 egg. Our results indicate that predator type and behavior can influence female behavior and nest fate, and that management actions that reduce the effectiveness of raccoons and skunks encountering waterfowl nests may benefit these nesting populations.

Journal of Wildlife Management↗

Occupancy Patterns of Breeding American Black Ducks

Occupancy patterns can assist with the determination of habitat limitation during breeding or wintering periods and can help guide population and habitat management efforts. American black ducks ( Anas rubripes ; black ducks) are thought to be limited by habitat and food availability during the winter, but breeding sites may also limit the size or growth potential of the population. The Canadian Wildlife Service conducts an annual breeding waterfowl survey that we used to explore the hypothesis that black duck carrying capacity is limited by wetlands available for breeding in Québec, Canada. We applied single‐visit, multi‐species occupancy models to the 1990–2015 population survey data to determine if there was evidence the black duck population was limited by breeding habitat. Using a dynamic (multi‐season) occupancy modeling approach, we estimated latent occupancy (occupancy accounting for imperfect detection) of black ducks and then used latent occupancy estimates to derive occupancy, colonization, and extirpation rates. We jointly modeled the occupancy dynamics of black ducks and other duck species in wetlands where both species were present. Throughout the duration of the survey, 44% of wetlands were never observed to be occupied by black ducks. Occupancy models showed wetland size was positively associated with occupancy at the first time step (initial occupancy) and colonization. All 2‐species models indicated initial black duck occupancy, persistence (continued occupancy), and colonization were positively associated with the presence of a second species. Colonization rate over the 26‐year period ranged from 7% to 27% across all models. Extirpation rates were similar and were constant through time within each model. Low occupancy rates, combined with approximately equal colonization and extirpation rates, suggest there are available wetlands for breeding black ducks in their core breeding area. If breeding habitats are not saturated, this suggests migration or wintering areas may be more limiting to black duck population abundance.

Ontario, Quebec↗

Body condition of wintering Pacific greater white-fronted geese

Extreme changes to key waterfowl habitats in the Klamath Basin (KB) on the Oregon–California border and the Sacramento Valley (SV) in California, USA, have occurred since 1980. The spatial distribution of Pacific greater white‐fronted geese ( Anser albifrons sponsa ; geese) has likewise changed among these areas and population size has grown from 79,000 to >600,000 geese during the same period. To assess the effects of landscape changes and spatial‐temporal distribution of geese, we collected Pacific greater white‐fronted geese during winters of 2009–2010 and 2010–2011 in the KB and SV and compared their body condition to geese collected during 1979–1980 and 1980–1981. We modeled body and lipid mass to assess body condition for each sex independently and examined the influence of collection day, year, and region. Body condition of geese varied throughout the winter and within years in a nonlinear fashion. We detected an increase in body condition in both sexes during December and January in the SV, which corresponds with improved habitat conditions and increases seen in other species in the region. Body condition upon arrival in fall migration varied by year for females and by year and region for males. Males and females arrived in poorer body condition during 2010–2011 than all other study years and males in the KB during 2010–2011 had extremely low lipid mass, reflecting poor regional habitat conditions induced by drought. Body condition of females varied over spring, by year, and by region and regional effects were evident for males. Body condition was significantly higher for geese in the SV than in the KB during spring. Our results suggest that Pacific greater white‐fronted geese have adapted to a changing landscape and have adjusted historical spatial use patterns to take advantage of more favorable conditions in the SV between 1979 and 2010.

California, Oregon↗

Great Lakes mallard population dynamics

Breeding mallard ( Anas platyrhynchos ) populations in the Great Lakes region (Michigan, Minnesota, Wisconsin, USA) declined by >40% between 2000–2022 based on abundance data collected during spring aerial surveys. Mallards are an important waterfowl species in this region, where an estimated 60–80% of the mallard harvest is composed of locally banded birds. Extensive population monitoring datasets are available for mallards, presenting an opportunity to address complex questions such as estimating productivity at large spatial and temporal scales, identifying the effects of harvest on mallard demography, quantifying mechanisms for harvest compensation, and integrating multiple datasets to quantify the demographic drivers of population change. Our objective was to simultaneously examine factors affecting demographic parameters and their relative contribution to Great Lakes mallard population dynamics. We used 32 years of banding, band recovery, and aerial survey data collected for mallards from Michigan and Wisconsin to develop an integrated population model (IPM). We used age ratios at banding to estimate productivity, band recoveries from hunter-harvested birds to estimate annual survival and cause-specific mortality (i.e., harvest or non-hunting), and modeled abundance using aerial survey and demographic parameter estimates from 1991–2022. The IPM results indicated the decline in Great Lakes mallard abundance was caused by increased non-hunting mortality and a decline in productivity. Productivity varied spatially but temporally declined with the loss of Conservation Reserve Program area. Moreover, our productivity assessment provided evidence of density dependence in reproduction. Non-hunting mortality was 3.5–6.7 times and 1.3–4.2 times greater than harvest mortality for adult and juvenile female mallards, respectively, indicating environmental factors during spring and summer, not harvest, most greatly influenced annual mortality for female mallards. Our IPM reduced uncertainty in the factors affecting Great Lakes mallard population dynamics and indicated management actions that address non-hunting mortality and productivity would be most effective in increasing Great Lakes mallard abundance.

Michigan, Wisconsin↗

Climate change, uncertainty, and natural resource management

Climate change and its associated uncertainties are of concern to natural resource managers. Although aspects of climate change may be novel (e.g., system change and nonstationarity), natural resource managers have long dealt with uncertainties and have developed corresponding approaches to decision-making. Adaptive resource management is an application of structured decision-making for recurrent decision problems with uncertainty, focusing on management objectives, and the reduction of uncertainty over time. We identified 4 types of uncertainty that characterize problems in natural resource management. We examined ways in which climate change is expected to exacerbate these uncertainties, as well as potential approaches to dealing with them. As a case study, we examined North American waterfowl harvest management and considered problems anticipated to result from climate change and potential solutions. Despite challenges expected to accompany the use of adaptive resource management to address problems associated with climate change, we conclude that adaptive resource management approaches will be the methods of choice for managers trying to deal with the uncertainties of climate change. ?? 2010 The Wildlife Society.

Journal of Wildlife Management↗

Harvest and dynamics of duck populations

The role of harvest in the dynamics of waterfowl populations continues to be debated among scientists and managers. Our perception is that interested members of the public and some managers believe that harvest influences North American duck populations based on calls for more conservative harvest regulations. A recent review of harvest and population dynamics of North American mallard (Anas platyrhynchos) populations (Pöysä et al. 2004) reached similar conclusions. Because of the importance of this issue, we reviewed the evidence for an impact of harvest on duck populations. Our understanding of the effects of harvest is limited because harvest effects are typically confounded with those of population density; regulations are typically most liberal when populations are greatest. This problem also exists in the current Adaptive Harvest Management Program (Conn and Kendall 2004). Consequently, even where harvest appears additive to other mortality, this may be an artifact of ignoring effects of population density. Overall, we found no compelling evidence for strong additive effects of harvest on survival in duck populations that could not be explained by other factors.

Journal of Wildlife Management↗

Stopover habitats of spring migrating surf scoters in southeast Alaska

Habitat conditions and nutrient reserve levels during spring migration have been suggested as important factors affecting population declines in waterfowl, emphasizing the need to identify key sites used during spring and understand habitat features and resource availability at stopover sites. We used satellite telemetry to identify stopover sites used by surf scoters migrating through southeast Alaska during spring. We then contrasted habitat features of these sites to those of random sites to determine habitat attributes corresponding to use by migrating scoters. We identified 14 stopover sites based on use by satellite tagged surf scoters from several wintering sites. We identified Lynn Canal as a particularly important stopover site for surf scoters originating throughout the Pacific winter range; approximately half of tagged coastally migrating surf scoters used this site, many for extended periods. Stopover sites were farther from the mainland coast and closer to herring spawn sites than random sites, whereas physical shoreline habitat attributes were generally poor predictors of site use. The geography and resource availability within southeast Alaska provides unique and potentially critical stopover habitat for spring migrating surf scoters. Our work identifies specific sites and habitat resources that deserve conservation and management consideration. Aggregations of birds are vulnerable to human activity impacts such as contaminant spills and resource management decisions. This information is of value to agencies and organizations responsible for emergency response planning, herring fisheries management, and bird and ecosystem conservation.

Journal of Wildlife Management↗

Band reporting probablilities of mallards, American black ducks, and wood ducks in eastern North America

Estimates of band reporting probabilities are used for managing North American waterfowl to convert band recovery probabilities into harvest probabilities, which are used to set harvest regulations. Band reporting probability is the probability that someone who has shot and retrieved a banded bird will report the band. This probability can vary relative to a number of factors, particularly the inscription on the band and the ease with which it can be reported. Other factors, such as geographic reporting region, and species and sex of the bird may also play a role. We tested whether reporting probabilities of wood ducks ( Aix sponsa ) and American black ducks (black ducks; Anas rubripes ) differed from those of mallards ( Anas platyrhynchos ) and whether band reporting varied geographically or by the sex of the banded bird. In the analysis of spatially comparable wood duck and mallard data, a band reporting probability of 0.73 (95% CI = 0.67–0.78) was appropriate for use across species, sex, and reporting region within the United States. In the black duck–mallard comparison, the band reporting probability of black ducks in Eastern Canada (0.50, 95% CI = 0.44–0.57) was lower than in the Eastern United States (0.73, 95% CI = 0.62–0.83). These estimates reflected an increase in overall band reporting probability following the addition of a toll-free telephone number to band inscriptions. Lower reporting in Eastern Canada may be because of cultural, linguistic, or logistical barriers.

Journal of Wildlife Management↗

Greater white-fronted goose habitat use in Louisiana provides water depth management insights

Numerous waterfowl species have altered their geographic distribution in recent decades. The greater white-fronted goose ( Anser albifrons ) has shifted its wintering distribution from coastal marshes in Texas and Louisiana, USA, to interior landscapes, creating challenges for conservation managers. Although the range shift has been primarily attributed to landscape-scale changes in rice production, effects of fine-scale habitat conditions on greater white-fronted goose habitat use are unknown. Thus, information on fine-scale environmental conditions could provide information to managers working to mitigate effects of the shift. We used global positioning system neck collars to monitor 78 greater white-fronted geese in Louisiana each winter during 2016–2019 and visited all used locations to record vegetation height, water depth, and land cover. We modeled greater white-fronted goose habitat use in 4 regions across Louisiana (central, coastal, north, southwest), and evaluated effects of vegetation height, water depth, land cover, and interactions. In our candidate model set, we included a sampling null, which accounted for differences in sampling frequencies among hunt seasons and diel periods. The top model in the central, coastal, and north regions included water depth, whereas the top model in the southwest region included water depth, vegetation height, and land cover. Parameter estimates and confidence intervals indicated that goose habitat use was higher at sites with >5.1 cm of water compared to dry sites or sites with <5.1 cm of water. In the southwest region, goose habitat use was highest in areas identified as other land cover (e.g., corn, cotton, rye, sugarcane) and lowest in open water. Although Louisiana has substantial anthropogenic food resources in the form of rice agriculture and abundant grazing opportunities afforded by mild winters, our results indicate that wet areas provide better habitat to greater white-fronted geese than dry areas. New agricultural flooding regimes in Louisiana could improve habitat quality and potentially mitigate some effects of the winter distribution shift.

Louisiana↗

Seasonal and decadal changes in winter body condition of four sympatric diving ducks

Winter body condition is an important driver of survival, reproductive output, and overall population health in waterfowl. Diving duck species use distinct habitats, exploit unique resources, and can collectively provide an integrated index of winter habitat quality. The San Francisco Bay (SFB) is the largest estuary on the west coast of North America, serving as critical wintering habitat for diving ducks in the Pacific Flyway. To evaluate the body condition of diving ducks after multiple decades of tidal habitat restoration in SFB, we compared structurally corrected measurements (annual and decadal) of total carcass lipid among canvasback ( Aythya valisineria ), greater scaup ( Aythya marila ), lesser scaup ( Aythya affinis ), and ruddy duck ( Oxyura jamaicensis ). We found greater and lesser scaup gained lipid reserves within the estuary prior to spring migration; however, canvasback and ruddy duck lipid levels declined throughout the winter and into the spring. Lipid loss over the wintering period could indicate a reduction in foraging resource availability, dietary shifts, or metabolic changes. These deficiencies could have negative cross-seasonal implications, as these species must accumulate resources outside of their wintering areas to fuel migration and reproduction. Lipid levels were elevated in lesser scaup collected in 2017-2019 compared to 1998-2000, while we found no decadal differences in lipid levels for canvasback or greater scaup. Our findings suggest tidal restoration in SFB, among other factors, may benefit some diving duck species.

California↗

Survival of surf scoters and white-winged scoters during remigial molt

Quantifying sources and timing of variation in demographic rates is necessary to determine where and when constraints may exist within the annual cycle of organisms. Surf scoters ( Melanitta perspicillata ) and white-winged scoters ( M. fusca ) undergo simultaneous remigial molt during which they are flightless for >1 month. Molt could result in reduced survival due to increased predation risk or increased energetic demands associated with regrowing flight feathers. Waterfowl survival during remigial molt varies across species, and has rarely been assessed for sea ducks. To quantify survival during remigial molt, we deployed very high frequency (VHF) transmitters on surf scoters ( n &thinsp;=&thinsp;108) and white-winged scoters ( n &thinsp;=&thinsp;57) in southeast Alaska and the Salish Sea (British Columbia and Washington) in 2008 and 2009. After censoring mortalities potentially related to capture and handling effects, we detected no mortalities during remigial molt; thus, estimates of daily and period survival for both scoter species during molt were 1.00. We performed sensitivity analyses in which mortalities were added to the dataset to simulate potential mortality rates for the population and then estimated the probability of obtaining a dataset with 0 mortalities. We found that only at high survival rates was there a high probability of observing 0 mortalities. We conclude that remigial molt is normally a period of low mortality in the annual cycle of scoters. The molt period does not appear to be a constraint on scoter populations; therefore, other annual cycle stages should be targeted by research and management efforts to change population trajectories.

Alaska, British Columbia, Washington↗

Reliability of external characteristics to age Barrow’s goldeneye

Accurate assignment of age class is critical for understanding most demographic processes. For waterfowl, most techniques for determining age class require birds in hand, reducing utility for quickly and efficiently sampling a large portion of the population. As an alternative, we sought to establish an observation‐based methodology, achievable in the field with standard optics, for determining age class of Barrow's goldeneyes ( Bucephala islandica ). We photographed heads, wings, and bellies of 232 Barrow's goldeneyes captured during late winter (February–April) of 2007–2015 along the north Pacific Coast. From these photographs, we focused on 5 external characteristics for both males and females, with binary states that putatively corresponded to 2 age classes—first‐year birds (<1 yr) and adults (>1 yr). For males, all 5 external traits (belly color, head color, eye color, facial crescent, median secondary coverts color) had binary states that were reliably distinguishable by observers. Moreover, all 5 external traits were highly predictive of age class (≥96% concordance between external vs. bursal‐derived ages), and novice observers, after receiving training, were able to accurately age 96% of first‐year and 99% of adult males. In contrast, patterns were weaker for females; putative external characteristics of female age class (belly color, bill radiance, bill blackness, eye color, median secondary coverts color) had 77–91% concordance with bursal‐derived age, compared with 96–100% for males, and observers misidentified age classes of 15% of females, compared with only 2% of males. Overall, age classes of male Barrow's goldeneyes were accurately and reliably distinguishable during winter based on several external characteristics, whereas those of females were not. Our technique may be used to estimate age composition of male Barrow's goldeneyes during winter, providing a useful metric for monitoring annual changes in adult‐to‐juvenile ratios and other important demographic parameters.

Alaska, British Columbia↗

Long-term spatial heterogeneity in mallard distribution in the Prairie pothole region

The Prairie Pothole Region (PPR) of north-central United States and south-central Canada supports greater than half of all breeding mallards ( Anas platyrhynchos ) annually counted in North America and is the focus of widespread conservation and research efforts. Allocation of conservation resources for this socioeconomically important population would benefit from an understanding of the nature of spatiotemporal variation in distribution of breeding mallards throughout the 850,000 km 2 landscape. We used mallard counts from the Waterfowl Breeding Population and Habitat Survey to test for spatial heterogeneity and identify high- and low-abundance regions of breeding mallards over a 50-year time series. We found strong annual spatial heterogeneity in all years: 90% of mallards counted annually were on an average of only 15% of surveyed segments. Using a local indicator of spatial autocorrelation, we found a relatively static distribution of low-count clusters in northern Montana, USA, and southern Alberta, Canada, and a dynamic distribution of high-count clusters throughout the study period. Distribution of high-count clusters shifted southeast from northwestern portions of the PPR in Alberta and western Saskatchewan, Canada, to North and South Dakota, USA, during the latter half of the study period. This spatial redistribution of core mallard breeding populations was likely driven by interactions between environmental variation that created favorable hydrological conditions for wetlands in the eastern PPR and dynamic land-use patterns related to upland cropping practices and government land-retirement programs. Our results highlight an opportunity for prioritizing relatively small regions within the PPR for allocation of wetland and grassland conservation for mallard populations. However, the extensive spatial heterogeneity in core distributions over our study period suggests such spatial prioritization will have to overcome challenges presented by dynamic land-use and climate patterns in the region, and thus merits additional monitoring and empirical research to anticipate future population distribution. Published 2017. This article is a U.S. Government work and is in the public domain in the USA.

Wildlife Society Bulletin↗

Communication strategies for reducing lead poisoning in wildlife and human health risks

Although lead poisoning in North American waterfowl has been reduced, it persists among other wildlife. To address this issue, we review lead poisoning in wildlife and threats to human health, describe the recent socio-political landscape, and develop a framework for reducing lead exposure related to hunting ammunition and fishing tackle. Despite substantial information about lead poisoning in wildlife, an explicit and strategic plan for using existing information to develop an effective communication program is lacking. Local and regional efforts encouraging hunters and anglers to voluntarily use nonlead alternatives could benefit from a nationally coordinated and strategic focus. We propose that the diffusion of innovation theory provides a useful framework for developing and implementing voluntary nonlead hunting ammunition and fishing tackle programs. Further, it can help communicators refine messages, increase efficiencies in developing communication materials, and monitor adoption of nonlead alternatives. The initial step in this process, however, is to engage stakeholders about the importance of the issue and leverage that concern as a catalyst for positive change. Published 2019. This article is a U.S. Government work and is in the public domain in the USA.

Wildlife Society Bulletin↗

Effects of fire on bird populations in mixed-grass prairie

The mixed-grass prairie is one of the largest ecosystems in North America, originally covering about 69 million ha (Bragg and Steuter 1995). Although much of the natural vegetation has been replaced by cropland and other uses (Samson and Knopf 1994, Bragg and Steuter 1995), significant areas have been preserved in national wildlife refuges, waterfowl production areas, state game management areas, and nature preserves. Mixed-grass prairie evolved with fire (Bragg 1995), and fire is frequently used as a management tool for prairie (Berkey et al. 1993).

Book chapter↗