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Environmental drivers of biseasonal anthrax outbreak dynamics in two multihost savanna systems

Environmental factors are common forces driving infectious disease dynamics. We compared interannual and seasonal patterns of anthrax infections in two multihost systems in southern Africa: Etosha National Park, Namibia, and Kruger National Park, South Africa. Using several decades of mortality data from each system, we assessed possible transmission mechanisms behind anthrax dynamics, examining (1) within- and between-species temporal case correlations and (2) associations between anthrax mortalities and environmental factors, specifically rainfall and the Normalized Difference Vegetation Index (NDVI), with empirical dynamic modeling. Anthrax cases in Kruger had wide interannual variation in case numbers, and large outbreaks seemed to follow a roughly decadal cycle. In contrast, outbreaks in Etosha were smaller in magnitude and occurred annually. In Etosha, the host species commonly affected remained consistent over several decades, although plains zebra ( Equus quagga ) became relatively more dominant. In Kruger, turnover of the main host species occurred after the 1990s, where the previously dominant host species, greater kudu ( Tragelaphus strepsiceros ), was replaced by impala ( Aepyceros melampus ). In both parks, anthrax infections showed two seasonal peaks, with each species having only one peak in a year. Zebra, springbok ( Antidorcas marsupialis ), wildebeest ( Connochaetes taurinus ), and impala cases peaked in wet seasons, while elephant ( Loxodonta africana ), kudu, and buffalo ( Syncerus caffer ) cases peaked in dry seasons. For common host species shared between the two parks, anthrax mortalities peaked in the same season in both systems. Among host species with cases peaking in the same season, anthrax mortalities were mostly synchronized, which implies similar transmission mechanisms or shared sources of exposure. Between seasons, outbreaks in one species may contribute to more cases in another species in the following season. Higher vegetation greenness was associated with more zebra and springbok anthrax mortalities in Etosha but fewer elephant cases in Kruger. These results suggest that host behavioral responses to changing environmental conditions may affect anthrax transmission risk, with differences in transmission mechanisms leading to multihost biseasonal outbreaks. This study reveals the dynamics and potential environmental drivers of anthrax in two savanna systems, providing a better understanding of factors driving biseasonal dynamics and outbreak variation among locations.

Etosha National Park, Kruger National Park

Climate change weakens the impact of disturbance interval on the growth rate of natural populations of Venus flytrap

Disturbances elicit both positive and negative effects on organisms; these effects vary in their strength and their timing. Effects of disturbance interval (i.e., the length of time between disturbances) on population growth will depend on both the timing and strength of positive and negative effects of disturbances. Climate change can modify the relative strengths of these positive and negative effects, leading to altered optimal disturbance intervals (the disturbance interval at which population growth rate is highest) and changes in the sensitivity of population growth rate to disturbance interval. While we know that climate may alter impacts of disturbance in some systems, we have a poor understanding of which effects of disturbance and which vital rates might drive an altered response to disturbance interval in a changing climate. We use demographic monitoring of natural populations of Dionaea muscipula , the Venus flytrap, that have experienced natural and managed fires, combined with realistic past and future climate projections, to construct climate- and fire-driven integral projection models (IPMs). We use these IPMs to compare the effect of fire return interval (FRI) on population growth rate in past and future climates. To dissect the mechanisms driving FRI response, we then construct IPMs with demographic data from an experimental manipulation of fire effects (ash addition, neighbor removal) and an accidental fire. Our results show that an FRI of 10 years is optimal for D. muscipula in past climate conditions, but a longer FRI (12 years) is optimal in future climate conditions. Further, deviations from optimal FRI reduce population growth rate dramatically in the past climate, but this reduction is muted in a future climate (future minus past sensitivity = 0.006, 95% CI [0.002, 0.011]). Finally, our experimental work suggests that fire effects are driven in part by positive, additive effects of competitor removal and ash addition immediately following a fire; for one population, both these treatments significantly increased population growth rate. Our work suggests that climate change can alter the response of populations to disturbance, highlighting the need to consider the interacting effects of multiple abiotic drivers when projecting future population growth and geographical distributions.

North Carolina

Tree symbioses sustain nitrogen fixation despite excess nitrogen supply

Symbiotic nitrogen fixation (SNF) is a key ecological process whose impact depends on the strategy of SNF regulation—the degree to which rates of SNF change in response to limitation by N versus other resources. SNF that is obligate or exhibits incomplete downregulation can result in excess N fixation, whereas a facultative SNF strategy does not. We hypothesized that tree-based SNF strategies differed by latitude (tropical vs. temperate) and symbiotic type (actinorhizal vs. rhizobial). Specifically, we expected tropical rhizobial symbioses to display strongly facultative SNF as an explanation of their success in low-latitude forests. In this study we used 15 N isotope dilution field experiments in New York, Oregon, and Hawaii to determine SNF strategies in six N-fixing tree symbioses. Nitrogen fertilization with +10 and +15 g N m −2 year −1 for 4–5 years alleviated N limitation in all taxa, paving the way to determine SNF strategies. Contrary to our hypothesis, all six of the symbioses we studied sustained SNF even at high N. Robinia pseudoacacia (temperate rhizobial) fixed 91% of its N (%N dfa ) in controls, compared to 64% and 59% in the +10 and +15 g N m −2 year −1 treatments. For Alnus rubra (temperate actinorhizal), %N dfa was 95%, 70%, and 60%. For the tropical species, %N dfa was 86%, 80%, and 82% for Gliricidia sepium (rhizobial); 79%, 69%, and 67% for Casuarina equisetifolia (actinorhizal); 91%, 42%, and 67% for Acacia koa (rhizobial); and 60%, 51%, and 19% for Morella faya (actinorhizal). Fertilization with phosphorus did not stimulate tree growth or SNF. These results suggest that the latitudinal abundance distribution of N-fixing trees is not caused by a shift in SNF strategy. They also help explain the excess N in many forests where N fixers are common.

Hawai'i, New York, Oregon

Hydrodynamics structure plankton communities and interactions in a freshwater tidal estuary

Drivers of phytoplankton and zooplankton dynamics vary spatially and temporally in estuaries due to variation in hydrodynamic exchange and residence time, complicating efforts to understand controls on food web productivity. We conducted approximately monthly (2012–2019; n = 74) longitudinal sampling at 10 fixed stations along a freshwater tidal terminal channel in the San Francisco Estuary, California, characterized by seaward to landward gradients in water residence time, turbidity, nutrient concentrations, and plankton community composition. We used multivariate autoregressive state space (MARSS) models to quantify environmental (abiotic) and biotic controls on phytoplankton and mesozooplankton biomass. The importance of specific abiotic drivers (e.g., water temperature, turbidity, nutrients) and trophic interactions differed significantly among hydrodynamic exchange zones with different mean residence times. Abiotic drivers explained more variation in phytoplankton and zooplankton dynamics than a model including only trophic interactions, but individual phytoplankton–zooplankton interactions explained more variation than individual abiotic drivers. Interactions between zooplankton and phytoplankton were strongest in landward reaches with the longest residence times and the highest zooplankton biomass. Interactions between cryptophytes and both copepods and cladocerans were stronger than interactions between bacillariophytes (diatoms) and zooplankton taxa, despite contributing less biovolume in all but the most landward reaches. Our results demonstrate that trophic interactions and their relative strengths vary in a hydrodynamic context, contributing to food web heterogeneity within estuaries at spatial scales smaller than the freshwater to marine transition.

California

Environmental variation structures reproduction and recruitment in long-lived mega-herbivores: Galapagos giant tortoises

Migratory, long-lived animals are an important focus for life-history theory because they manifest extreme trade-offs in life-history traits: delayed maturity, low fecundity, variable recruitment rates, long generation times, and vital rates that respond to variation across environments. Galapagos tortoises are an iconic example: they are long-lived, migrate seasonally, face multiple anthropogenic threats, and have cryptic early life-history stages for which vital rates are unknown. From 2012 to 2021, we studied the reproductive ecology of two species of Galapagos tortoises ( Chelonoidis porteri and C. donfaustoi ) along elevation gradients that coincided with substantial changes in climate and vegetation productivity. Specifically, we (1) measured the body and reproductive condition of 166 adult females, (2) tracked the movements of 33 adult females using global positioning system telemetry, and monitored their body condition seasonally, (3) recorded nest temperatures, clutch characteristics, and egg survival from 107 nests, and (4) used radiotelemetry to monitor growth, survival, and movements of 104 hatchlings. We also monitored temperature and rainfall from field sites, and remotely sensed primary productivity along the elevation gradient. Our study showed that environmental variability, mediated by elevation, influenced vital rates of giant tortoises, specifically egg production by adult females and juvenile recruitment. Adult females were either elevational migrants or year-round lowland residents. Migrants had higher body condition than residents, and body condition was positively correlated with the probability of being gravid. Nests occurred in the hottest, driest parts of the tortoise's range, between 6 and 165 m elevation. Clutch size increased with elevation, whereas egg survival decreased. Hatchling survival and growth were highest at intermediate elevations. Hatchlings dispersed rapidly to 100–750 m from their nests before becoming sedentary (ranging over <0.2 ha). Predicted future climates may impact the relationships between elevation and vital rates of Galapagos tortoises and other species living across elevation gradients. Resilience will be maximized by ensuring the connectivity of foraging and reproductive areas within the current and possible future elevational ranges of these species.

Galapagos Islands, Santa Cruz Island

Using mobile acoustic monitoring and false-positive N-mixture models to estimate bat abundance and population trends

Estimating the abundance of unmarked animal populations from acoustic data is challenging due to the inability to identify individuals and the need to adjust for observation biases including detectability (false negatives), species misclassification (false positives), and sampling exposure. Acoustic surveys conducted along mobile transects were designed to avoid counting individuals more than once, where raw counts are commonly treated as an index of abundance. More recently, false-positive abundance models have been developed to estimate abundance while accounting for imperfect detection and misclassification. We adapted these methods to model summertime abundance and trends of three species of bats at multiple spatial scales using acoustic recordings collected along mobile transects by partners of the North American Bat Monitoring Program (NABat) from 2012 to 2020. This multiscale modeling spanned individual transect routes, larger NABat grid cells (10 km × 10 km), and across the entire extent of modeled species ranges. We estimated relationships between species abundances and a suite of abiotic and biotic predictors (landcover types, climatological variables, physiographic diversity, building density, and the impacts of white-nose syndrome [WNS]) and found varying levels of support between species. We present clear evidence of substantial declines in populations of tricolored bats ( Perimyotis subflavus ) and little brown bats ( Myotis lucifugus ), declines that corresponded in space and time with the progression of WNS, a devastating disease of hibernating bats. In contrast, our analysis revealed that similar population-wide declines probably have not occurred in big brown bats ( Eptesicus fuscus ), a species known to be less affected by WNS. This study provides the first abundance-based species distribution predictions and population trends for bats in their summer ranges in North America. These models will probably be applicable to assessing wildlife populations in other monitoring programs where acoustic data are used or where false-negative and false-positive detections are present. Finally, our abundance framework (as a spatial point pattern process) can serve as a foundation from which more sophisticated integrated species distribution models that incorporate additional streams of monitoring data (e.g., stationary acoustics, captures) can be developed for North American bats.

Ecological Monographs

An integrative paradigm for building causal knowledge

A core aspiration of the ecological sciences is to determine how systems work, which implies the challenge of developing a causal understanding. Causal inference has long been approached from a statistical perspective, which can be limited and restrictive for a variety of reasons. Ecologists and other natural scientists have historically pursued mechanistic knowledge as an alternative approach to causal understanding, though without explicit reference to the requirements of causal statistics. In this paper, I describe the premises of an expanded paradigm for causal studies, the Integrative Causal Investigation Paradigm, that subsumes causal statistics and mechanistic investigation into a multi-evidence approach. This paradigm is distinct from the one articulated by causal statistics in that it (1) focuses its attention on the long-term goal of building causal knowledge across multiple studies and (2) recognizes the essential role of mechanistic investigations in establishing a causal understanding. The Integrative Paradigm, consequentially, proposes that there are multiple methodological routes to building causal knowledge and thus represents a pluralistic perspective. This paper begins by describing the crux of the problem faced by causal statistics. To understand this problem, it should be recognized that the word causal has multiple meanings and a variety of evidential standards. An expanded vocabulary is developed so as to reduce ambiguities and clarify critical issues. I further show by example that there is an important ingredient typically omitted from consideration in causal statistics, which is the known information related to the mechanisms underlying relationships being evaluated. To address this issue, I describe a procedure, Causal Knowledge Analysis, that involves an evaluation and compilation of existing evidence indicative of causal content and the features of mechanisms. Causal Knowledge Analysis is applied to three example situations to illustrate the process and its potential for contributing to the development of causal knowledge. The implications of adopting the proposed paradigm and associated procedures are discussed and include the potential for advancing ecology, the potential for clarifying causal methodology, and the potential for contributing to predictive forecasting.

Ecological Monographs

Hotter temperatures alter riparian plant outcomes under regulated river conditions

Climate change and river regulation alter environmental controls on riparian plant occurrence and cover worldwide. Simultaneous changes to river flow and air temperature could result in unanticipated plant responses to novel environmental conditions. Increasing temperature could alter riparian plant response to hydrology and other factors, while river regulation may exacerbate environmental stress through novel flows like those resulting from power generation. Further, plant establishment and growth may require differing conditions, which may be decoupled by novel conditions. Using a large dataset that spans a natural 5°C mean annual temperature (MAT) gradient and a Bayesian model that integrates plant occurrence and cover, we address four questions: (1) Does hotter MAT modify plant response to hydrology, substrate composition, topography, and cover of co-occurring plant species? (2) Does the timing of hydropower tides benefit some species over others? (3) Does dam-induced erosion hinder riparian species more than upland species? (4) Do occurrence and cover respond to different environmental variables, allowing for decoupling of life history processes? We addressed these questions with data collected along 364 km of the Colorado River downstream of Glen Canyon Dam, Arizona, United States of America. Occurrence and cover class were recorded in >10,000 plots from 2016 to 2020, along with environmental covariates that repeat across the climate gradient. For 36 species, plant occurrence and cover were modeled with respect to MAT, hydrology, substrate, topography, other plant cover, and their interactions with MAT. There were four key results. (1) Increasing MAT will not only directly influence plants but will mediate their responses to the environment, including greater dependence on stable water supplies. (2) The timing of hydropower tides shapes plant community composition. (3) Dam-related erosion has an outsized effect on riparian species, which could lead to a loss of regionally unique plant species. (4) For all species, the most important covariates driving occurrence differed from those for cover, suggesting the potential for these life stages to be decoupled. Not only will climate change and river regulation independently alter plant distributions, interactions among hotter temperature, dam-controlled flow patterns, and limited fine sediments will determine which species flourish or perish under future conditions.

Arizona

Mammalian predator co‐occurrence affected by prey and habitat more than competitor presence at multiple time scales

The behavior and abundance of sympatric predators can be affected by a complex dominance hierarchy. The strength of antagonistic interactions in predator communities is difficult to study and remains poorly understood for many predator assemblages. Predators directly and indirectly influence the broader ecosystem, so identifying the relative importance of competition, prey, and habitat in shaping predator interactions has broad conservation and management implications. We investigated space use among five predator species (black bear [ Ursus americanus ], bobcat [ Lynx rufus ], coyote [ Canis latrans ], mountain lion [ Puma concolor ], and gray wolf [ Canis lupus ]) across three temporal scales in northern Idaho, USA. We used camera trap data to test whether potentially subordinate predators spatially avoided dominant predators and how prey availability influenced those relationships. We found few instances of subordinate predators spatially avoiding dominant predators and only at the finest temporal scale of our analyses. Instead, habitat features generally influenced predator space use patterns at coarser scales whereas prey and competitor presence influenced space use patterns at finer scales. Co-occurrence was positively associated between coyotes and bobcats at coarser timescales and between mesopredators and apex predators at finer timescales. Bobcats and mountain lions temporarily delayed the use of sites recently visited by coyotes and black bears, respectively. And all predator species used sites sooner following the detection of a competitor in areas with higher relative abundances of prey (primarily white-tailed deer [ Odocoileus virginianus ]). Our results suggest attraction to shared habitats and prey resources influenced space use in the predator community more than avoidance of competitors. We propose that the effects of interspecific interactions on predator distributions were most evident for mesopredators because their trophic position requires balancing risks and rewards associated with prey, apex predators, and other mesopredators. In addition, relatively high densities of a common prey source likely facilitated the spatial coexistence in this predator community. Our study demonstrates the value of simultaneously assessing multiple interspecific interactions across different spatiotemporal scales to discern relationships within the predator guild.

Idaho

Pan-amphibia distribution of the fungal parasite Batrachochytrium dendrobatidis varies with species and temperature

Batrachochytrium dendrobatidis (Bd) is a globally distributed fungal pathogen of amphibians that has contributed to one of the largest disease-related biodiversity losses in wildlife. Bd is regularly viewed through the lens of a global wildlife epizootic because the spread of highly virulent genetic lineages has resulted in well-documented declines and extinctions of multiple amphibian species. However, the current state of Bd occurrence, host range, host impacts, and ecological drivers remains poorly understood outside of the most negatively affected amphibian species and regions. Our objective was to describe the macroecology of Bd occurrence and infection intensity on caudates (salamanders) across the United States and to compare these patterns with better-studied anurans (frogs and toads). We collected swabs from 11,183 amphibians at 609 sites from 54 species across the United States from 2015 to 2017. We analyzed the prevalence and intensity of Bd infection jointly using a Bayesian hurdle model with covariates of site-level temperature and precipitation, as well as individual characteristics and species identification. Bd was distributed widely across sites and species sampled across the spatial extent of the conterminous United States. We found that Bd prevalence and intensity were most strongly influenced by temperature in the month preceding sampling and by differences among taxon groups. We estimated that temperature had a strong and nonlinear influence on both Bd prevalence and intensity with peak infection at intermediate temperatures and lower infection at low and high temperatures. We found Caudate hosts tended to have higher prevalence than Anuran hosts and Anuran hosts tended to have higher intensity at optimal temperatures for Bd infection. Our findings suggest that Bd has an amphibian-wide host range, temperature gradients exert a strong influence on Bd, and enzootic transmission likely encompasses a much larger spatial and species distribution than previously recognized across North America.

conterminous United States

Missing data in ecology: Syntheses, clarifications, and considerations

In ecology and related sciences, missing data are common and occur in a variety of different contexts. When missing data are not handled properly, subsequent statistical estimates tend to be biased, inefficient, and lack proper confidence interval coverage. Missing data are often grouped into three categories: missing completely at random (MCAR), missing at random (MAR), and missing not at random (MNAR). We review each category and compare their benefits and drawbacks. We review several approaches to handling missing data including complete case analysis, imputation, inverse probability weighting, and data augmentation. We clarify what types of variables should accompany imputation methods and how those variables are influenced by the analysis methods. Additionally, we discuss missing data that lack a formal basis for measurement and hence are fundamentally different from MCAR, MAR, and MNAR missing data. Throughout, we introduce concepts and numeric examples using both simulated data and data from the United States Environmental Protection Agency's 2016 National Wetland Condition Assessment. We conclude by providing five considerations for ecologists and other scientists handling missing data.

Ecological Monographs

Geographic distribution of the mid-continent population of sandhill cranes and related management applications

The Mid-continent Population (MCP) of sandhill cranes ( Grus canadensis ) is widely hunted in North America and is separated into the Gulf Coast Subpopulation and Western Subpopulation for management purposes. Effective harvest management of the MCP requires detailed knowledge of breeding distribution of subspecies and subpopulations, chronology of their use of fall staging areas and wintering grounds, and exposure to and harvest from hunting. To address these information needs, we tagged 153 sandhill cranes with Platform Transmitting Terminals (PTTs) during 22 February–12 April 1998–2003 in the Central and North Platte River valleys of south-central Nebraska. We monitored PTT-tagged sandhill cranes, hereafter tagged cranes, from their arrival to departure from breeding grounds, during their fall migration, and throughout winter using the Argos satellite tracking system. The tracking effort yielded 74,041 useable locations over 49,350 tag days; median duration of tracking of individual cranes was 352 days and 73 cranes were tracked >12 months. Genetic sequencing of mitochondrial DNA (mtDNA) from blood samples taken from each of our random sample of tagged cranes indicated 64% were G. c. canadensis and 34% were Grus canadensis tabida . Tagged cranes during the breeding season settled in northern temperate, subarctic, and arctic North America (U.S. [23%, n = 35], Canada [57%, n = 87]) and arctic regions of northeast Asia (Russia [20%, n = 31]). Distribution of tagged cranes by breeding affiliation was as follows: Western Alaska–Siberia (WA–S, 42 ± 4% [SE]), northern Canada–Nunavut (NC–N, 21 ± 4%), west-central Canada–Alaska (WC–A, 23 ± 4%) and East-central Canada–Minnesota (EC–M, 14 ± 3%). All tagged cranes returned to the same breeding affiliation used during the previous year with a median distance of 1.60 km (range: 0.08–7.7 km, n = 53) separating sites used in year 1 and year 2. Fall staging occurred primarily in central and western Saskatchewan (69%), North Dakota (16%), southwestern Manitoba (10%), and northwestern Minnesota (3%). Space-use sharing indices showed that except for NC–N and WC–A birds, probability of finding a crane from one breeding affiliation within the home range of another breeding affiliation was low during fall staging. Tagged cranes from WC–A and EC–M breeding affiliations, on average, spent 25 and 20 days, respectively, longer on fall staging areas in the northern plains than did WA–S and NC–N birds. Cranes in the NC–N, WA–S, and WC–A affiliations spent 99%, 74%, and 64%, respectively, of winter in western Texas in Hunting Zone A; EC–M cranes spent 83% of winter along the Texas Gulf Coast in Hunting Zone C. Tagged cranes that settled within the breeding range of the Gulf Coast Subpopulation spent 28% and 42% of fall staging and winter within the range of the Western Subpopulation, indicating sufficient exchange of birds to potentially limit effectiveness of MCP harvest management. Harvests of EC–M and WC–A cranes during 1998–2003 were disproportionately high to their estimated numbers in the MCP, suggesting more conservative harvest strategies may be required for these subpopulations in the future, and for sandhill cranes to occupy major parts of their historical breeding range in the Prairie Pothole Region. Exceptionally high philopatry of MCP cranes of all 4 subpopulations to breeding sites coupled with strong linkages between crane breeding distribution, and fall staging areas and wintering grounds, provide managers guidance for targeting MCP crane harvest to meet management goals. Sufficient temporal or spatial separation exists among the 4 subpopulations on fall staging areas and wintering grounds to allow harvest to be targeted at the subpopulation level in all states and provinces (and most hunting zones within states and provinces) when conditions warrant. Knowledge gained from our study provides decision-makers in the United States, Canada, Mexico, and Russia with improved guidance for developing sound harvest regulations, focusing conservation efforts, and generating collaborative efforts among these nations on sandhill crane research and management to meet mutually important goals.

Nebraska

Competitive interactions and resource partitioning between northern spotted owls and barred owls in western Oregon

The federally threatened northern spotted owl ( Strix occidentalis caurina ) is the focus of intensive conservation efforts that have led to much forested land being reserved as habitat for the owl and associated wildlife species throughout the Pacific Northwest of the United States. Recently, however, a relatively new threat to spotted owls has emerged in the form of an invasive competitor: the congeneric barred owl ( S. varia ). As barred owls have rapidly expanded their populations into the entire range of the northern spotted owl, mounting evidence indicates that they are displacing, hybridizing with, and even killing spotted owls. The range expansion by barred owls into western North America has made an already complex conservation issue even more contentious, and a lack of information on the ecological relationships between the 2 species has hampered recovery efforts for northern spotted owls. We investigated spatial relationships, habitat use, diets, survival, and reproduction of sympatric spotted owls and barred owls in western Oregon, USA, during 2007–2009. Our overall objective was to determine the potential for and possible consequences of competition for space, habitat, and food between these previously allopatric owl species. Our study included 29 spotted owls and 28 barred owls that were radio-marked in 36 neighboring territories and monitored over a 24-month period. Based on repeated surveys of both species, the number of territories occupied by pairs of barred owls in the 745-km 2 study area (82) greatly outnumbered those occupied by pairs of spotted owls (15). Estimates of mean size of home ranges and core-use areas of spotted owls (1,843 ha and 305 ha, respectively) were 2–4 times larger than those of barred owls (581 ha and 188 ha, respectively). Individual spotted and barred owls in adjacent territories often had overlapping home ranges, but interspecific space sharing was largely restricted to broader foraging areas in the home range with minimal spatial overlap among core-use areas. We used an information-theoretic approach to rank discrete-choice models representing alternative hypotheses about the influence of forest conditions, topography, and interspecific interactions on species-specific patterns of nighttime resource selection. Spotted owls spent a disproportionate amount of time foraging on steep slopes in ravines dominated by old (>120 yr) conifer trees. Barred owls used available forest types more evenly than spotted owls, and were most strongly associated with patches of large hardwood and conifer trees that occupied relatively flat areas along streams. Spotted and barred owls differed in the relative use of old conifer forest (greater for spotted owls) and slope conditions (steeper slopes for spotted owls), but we found no evidence that the 2 species differed in their use of young, mature, and riparian-hardwood forest types. Mean overlap in proportional use of different forest types between individual spotted owls and barred owls in adjacent territories was 81% (range = 30–99%). The best model of habitat use for spotted owls indicated that the relative probability of a location being used was substantially reduced if the location was within or in close proximity to a core-use area of a barred owl. We used pellet analysis and measures of food-niche overlap to determine the potential for dietary competition between spatially associated pairs of spotted owls and barred owls. We identified 1,223 prey items from 15 territories occupied by spotted owls and 4,299 prey items from 24 territories occupied by barred owls. Diets of both species were dominated by nocturnal mammals, but diets of barred owls included many terrestrial, aquatic, and diurnal prey species that were rare or absent in diets of spotted owls. Northern flying squirrels ( Glaucomys sabrinus ), woodrats ( Neotoma fuscipes , N. cinerea ), and lagomorphs ( Lepus americanus , Sylvilagus bachmani ) were primary prey for both owl species, accounting for 81% and 49% of total dietary biomass for spotted owls and barred owls, respectively. Mean dietary overlap between pairs of spotted and barred owls in adjacent territories was moderate (42%; range = 28–70%). Barred owls displayed demographic superiority over spotted owls; annual survival probability of spotted owls from known-fate analyses (0.81, SE = 0.05) was lower than that of barred owls (0.92, SE = 0.04), and pairs of barred owls produced an average of 4.4 times more young than pairs of spotted owls over a 3-year period. We found a strong, positive relationship between seasonal (6-month) survival probabilities of both species and the proportion of old (>120 yr) conifer forest within individual home ranges, which suggested that availability of old forest was a potential limiting factor in the competitive relationship between these 2 species. The annual number of young produced by spotted owls increased linearly with increasing distance from a territory center of a pair of barred owls, and all spotted owls that attempted to nest within 1.5 km of a nest used by barred owls failed to successfully produce young. We identified strong associations between the presence of barred owls and the behavior and fitness potential of spotted owls, as shown by changes in movements, habitat use, and reproductive output of spotted owls exposed to different levels of spatial overlap with territorial barred owls. When viewed collectively, our results support the hypothesis that interference competition with barred owls for territorial space can constrain the availability of critical resources required for successful recruitment and reproduction of spotted owls. Availability of old forests and associated prey species appeared to be the most strongly limiting factors in the competitive relationship between these species, indicating that further loss of these conditions can lead to increases in competitive pressure. Our findings have broad implications for the conservation of spotted owls, as they suggest that spatial heterogeneity in vital rates may not arise solely because of differences among territories in the quality or abundance of forest habitat, but also because of the spatial distribution of a newly established competitor. Experimental removal of barred owls could be used to test this hypothesis and determine whether localized control of barred owl numbers is an ecologically practical and socio-politically acceptable management tool to consider in conservation strategies for spotted owls.

Oregon

Spring migration ecology of the mid-continent sandhill crane population with an emphasis on use of the Central Platte River Valley, Nebraska

We conducted a 10-year study (1998–2007) of the Mid-Continent Population (MCP) of sandhill cranes (Grus canadensis) to identify spring-migration corridors, locations of major stopovers, and migration chronology by crane breeding affiliation (western Alaska–Siberia [WA–S], northern Canada–Nunavut [NC–N], west-central Canada–Alaska [WC–A], and east-central Canada–Minnesota [EC–M]). In the Central Platte River Valley (CPRV) of Nebraska, we evaluated factors influencing staging chronology, food habits, fat storage, and habitat use of sandhill cranes. We compared our findings to results from the Platte River Ecology Study conducted during 1978–1980. We determined spring migration corridors used by the breeding affiliations (designated subpopulations for management purposes) by monitoring 169 cranes marked with platform transmitter terminals (PTTs). We also marked and monitored 456 cranes in the CPRV with very high frequency (VHF) transmitters to evaluate length and pattern of stay, habitat use, and movements. An estimated 42% and 58% of cranes staging in the CPRV were greater sandhill cranes (G. c. tabida) and lesser sandhill cranes (G. c. canadensis), and they stayed for an average of 20 and 25 days (2000–2007), respectively. Cranes from the WA–S, NC–N, WC–A, and EC–M affiliations spent an average of 72, 77, 52, and 53 days, respectively, in spring migration of which 28, 23, 24, and 18 days occurred in the CPRV. The majority of the WA–S subpopulation settled in the CPRV apparently because of inadequate habitat to support more birds upstream, although WA–S cranes accounted for >90% of birds staging in the North Platte River Valley. Crane staging duration in the CPRV was negatively correlated with arrival dates; 92% of cranes stayed >7 days. A program of annual mechanical removal of mature stands of woody growth and seedlings that began in the early 1980s primarily in the main channel of the Platte River has allowed distribution of crane roosts to remain relatively stable over the past 2 decades. Most cranes returned to nocturnal roost sites used in previous years. Corn residues dominated the diet of sandhill cranes in the CPRV, as in the 1970s, despite a marked decline in standing crop of corn residues. Only 14% (10 of 74) of PTT-marked migrant cranes stayed at stopovers for ≥5 days before arriving in the CPRV, which limited the contribution of sites south of the CPRV for fat accumulation needed for migration and reproduction. Body masses of cranes (after adjusting for body size [an index of fat]) at arrival in the CPRV varied widely among years (1998–2006), indicating the importance of maintaining productive habitats on the wintering grounds to condition cranes for migration and reproduction. Average rates of fat gain by adult females while in the CPRV remained similar from 1978–1979 to 1998–1999 but declined among males. Distances cranes flew to feeding grounds in the CPRV increased as the percentage of cropland planted to soybeans increased and as density of cranes on nocturnal roosts increased. These results suggest that as habitats of limited or no value to cranes increase on the landscape, more flight time and higher maintenance costs may reduce fat storage. An estimated 40% of diurnal use occurred north of Interstate 80 (I-80) where ≤5% of lands dedicated to crane conservation are located. Seventy-four and 40% of PTT-marked EC–M and WC–A cranes had spring migrations that included staging in eastern South Dakota for an average of 11 and 10 days, respectively. Cranes of the NC–N, WA–S, and WC–A subpopulations staged an average of 25, 17, and 12 days in central and western Saskatchewan/eastern Alberta. Females in these affiliations increased their fat reserves after leaving Nebraska by an estimated 450, 451, and 452 g, respectively, underscoring the key role of these staging areas in preparing the 3 subpopulations for reproduction. After departing Nebraska, MCP cranes roosted primarily in basin wetlands. Most of these wetlands are in private ownership and lack adequate protection, emphasizing the need for effective laws and policies to ensure their long-term protection. The continued success of the current management goal of maintaining the MCP at approximately its current size and providing diverse recreational opportunities over a wide area of midcontinent and western North America is predicated on the ability of MCP cranes to continue to store large fat reserves in the CPRV in advance of breeding. For the CPRV to remain a key fat storage site, active channel maintenance (e.g., clearing of woody vegetation) likely will need to continue, along with establishing minimum stream flows. These actions would help ensure nocturnal roosting habitat remains sufficiently dispersed to provide cranes with daily intake of high-energy food adequate for major fat storage and limit risk of high mortality from storms and disease. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.

Nebraska

Habitat prioritization across large landscapes, multiple seasons, and novel areas: an example using greater sage-grouse in Wyoming

Animal habitat selection is an important and expansive area of research in ecology. In particular, the study of habitat selection is critical in habitat prioritization efforts for species of conservation concern. Landscape planning for species is happening at ever-increasing extents because of the appreciation for the role of landscape-scale patterns in species persistence coupled to improved datasets for species and habitats, and the expanding and intensifying footprint of human land uses on the landscape. We present a large-scale collaborative effort to develop habitat selection models across large landscapes and multiple seasons for prioritizing habitat for a species of conservation concern. Greater sage-grouse ( Centrocercus urophasianus , hereafter sage-grouse) occur in western semi-arid landscapes in North America. Range-wide population declines of this species have been documented, and it is currently considered as “warranted but precluded” from listing under the United States Endangered Species Act. Wyoming is predicted to remain a stronghold for sage-grouse populations and contains approximately 37% of remaining birds. We compiled location data from 14 unique radiotelemetry studies (data collected 1994–2010) and habitat data from high-quality, biologically relevant, geographic information system (GIS) layers across Wyoming. We developed habitat selection models for greater sage-grouse across Wyoming for 3 distinct life stages: 1) nesting, 2) summer, and 3) winter. We developed patch and landscape models across 4 extents, producing statewide and regional (southwest, central, northeast) models for Wyoming. Habitat selection varied among regions and seasons, yet preferred habitat attributes generally matched the extensive literature on sage-grouse seasonal habitat requirements. Across seasons and regions, birds preferred areas with greater percentage sagebrush cover and avoided paved roads, agriculture, and forested areas. Birds consistently preferred areas with higher precipitation in the summer and avoided rugged terrain in the winter. Selection for sagebrush cover varied regionally with stronger selection in the Northeast region, likely because of limited availability, whereas avoidance of paved roads was fairly consistent across regions. We chose resource selection function (RSF) thresholds for each model set (seasonal × regional combination) that delineated important seasonal habitats for sage-grouse. Each model set showed good validation and discriminatory capabilities within study-site boundaries. We applied the nesting-season models to a novel area not included in model development. The percentage of independent nest locations that fell directly within identified important habitat was not overly impressive in the novel area (49%); however, including a 500-m buffer around important habitat captured 98% of independent nest locations within the novel area. We also used leks and associated peak male counts as a proxy for nesting habitat outside of the study sites used to develop the models. A 1.5-km buffer around the important nesting habitat boundaries included 77% of males counted at leks in Wyoming outside of the study sites. Data were not available to quantitatively test the performance of the summer and winter models outside our study sites. The collection of models presented here represents large-scale resource-management planning tools that are a significant advancement to previous tools in terms of spatial and temporal resolution.

Wyoming

Demographic rates and population viability of black bears in Louisiana

The Louisiana black bear ( Ursus americanus luteolus ) was reduced to a few small, fragmented, and isolated subpopulations in the Lower Mississippi Alluvial Valley by the mid-twentieth century resulting from loss and fragmentation of habitat. In 1992, the United States Fish and Wildlife Service (USFWS) granted the Louisiana black bear threatened status under the United States Endangered Species Act of 1973. Since that time, a recovery plan was developed, a reintroduced population was established, and habitat recovery has occurred. The Recovery Plan states that a minimum of 2 populations must be viable (i.e., persistence probabilities over 100 years >0.95), 1 in the Tensas River Basin and 1 in the Atchafalaya River Basin. Consequently, our objectives were to 1) estimate demographic rates of Louisiana black bear subpopulations, 2) develop data-driven stochastic population projection models, and 3) determine how different projection model assumptions affect population trajectories and predictions about long-term persistence. Our overall goal was to assess long-term persistence of the bear subpopulations in Louisiana, individually and as a whole. We collected data using varying combinations of non-invasive DNA sampling, live capture, winter den visits, and radio monitoring from 2002 to 2012 in the 4 areas currently supporting breeding subpopulations in Louisiana: Tensas River Basin (TRB), Upper Atchafalaya River Basin (UARB), Lower Atchafalaya River Basin (LARB), and a recently reintroduced population at the Three Rivers Complex (TRC). From 2002 to 2012, we radio monitored fates of 86 adult females within the TRB and 43 in the TRC. Mean estimates of annual adult survival for the TRB and TRC were 0.997 and 0.990, respectively, when unknown fates were assumed alive and 0.970 and 0.926 when unknown fates were assumed dead. From 2003 to 2013, we observed 130 cub litters from 74 females in the TRB, and 74 cub litters from 45 females in the TRC. During the same period, we observed 43 yearling litters for 33 females in the TRB and 21 yearling litters for 19 females in the TRC. The estimated number of cubs and number of yearlings produced per breeding adult female was 0.47 and 0.20, respectively, in the TRB and 0.32 and 0.18 in the TRC. On the basis of matrix projection models, asymptotic growth rates ranged from 1.053 to 1.078 for the TRB and from 1.005 to 1.062 for the TRC, depending on how we treated unresolved fates of adult females. Persistence probabilities estimated from stochastic population models based on telemetry data ranged from 0.997 to 0.998 for the TRC subpopulation depending on model assumptions and were >0.999 for the TRB regardless of model assumptions. We extracted DNA from hair collected at baited, barbed-wire enclosures in the TRB, UARB, and LARB to determine individual identities for capture-mark-recapture (CMR) analysis. We used those detection histories to estimate apparent survival ( φ ), per-capita recruitment ( f ), abundance ( N ), realized growth rate ( λ ), and long-term viability, based on Bayesian hierarchical modeling methods that allowed estimation of temporal process variance and parameter uncertainty. Based on 23,312 hair samples, annual N for females in the TRB ranged from 133 to 164 during 2006–2012, depending on year and how detection heterogeneity was modeled. Geometric mean of λ ranged from 0.996 to 1.002. In the UARB, we collected 11,643 hair samples from 2007 to 2012, from which estimates of N for females ranged from 23 to 43 during the study period, depending on detection heterogeneity model. The geometric mean of λ ranged from 1.038 to 1.059. Estimated N for females in LARB ranged from 69 to 96, and annual λ ranged from 0.80 to 1.11 based on 3,698 hair samples collected during 2010–2012, also depending on year and heterogeneity model. Probabilities of persistence over 100 years for the TRC and TRB based on stochastic matrix projection models that used vital rate estimates from telemetry data were >0.95 for all scenarios. Probability of persistence at the TRB and the UARB based on projection models that used vital rate estimates from CMR analyses ranged from 0.928 to 0.954 and from 0.906 to 0.959, respectively, depending on model assumptions. Data from the LARB were insufficient for a viability assessment. Thus, individual persistence probabilities for TRB and UARB did not meet the strict definition of viability (i.e., >0.95) under some model assumptions. However, the joint probability of bears persisting either in the TRB or UARB was >0.993 assuming individual population dynamics were independent and was >0.958 assuming dynamics were perfectly correlated. Furthermore, including the TRC increased the joint probability of bears persisting somewhere in the TRB, UARB, or TRC to >0.999 based on the most pessimistic individual persistence estimates from those subpopulations. Therefore, if the intent of specifying that 2 subpopulations should be viable was to ensure the persistence of Louisiana black bears somewhere within its historical range, then the viability threshold was met. © 2016 The Wildlife Society.

Louisiana

Biological and social outcomes of antler point restriction harvest regulations for white-tailed deer

Selective harvest criteria, such as antler point restrictions (APRs), have been used to regulate harvest of male ungulates; however, comprehensive evaluation of the biological and social responses to this management strategy is lacking. In 2002, Pennsylvania adopted new APRs for white-tailed deer ( Odocoileus virginianus ) that required, depending on wildlife management unit, ≥3 or ≥4 points on 1 antler for legal harvest. Historically, harvest rates of subadult (1.5 yr old) and adult (≥2.5 yr old) antlered males averaged 0.80. Antler point restrictions were designed to protect ≥50% of subadult males from harvest. Most adult males remained legal for harvest. We estimated harvest rates, survival rates, and cause-specific mortality of radio-collared male deer (453 subadults, 103 adults) in 2 wildlife management units (Armstrong and Centre counties) to evaluate biological efficacy of APRs to increase recruitment of adult males during 2002–2005. We administered statewide deer hunter surveys before and after each hunting season over the same 3 years to evaluate hunter attitudes toward APRs. We conducted 2 types of surveys: a simple random sample of all license buyers for each survey and a longitudinal panel of hunters who completed all 6 surveys. At the same time APRs were implemented, the Pennsylvania Game Commission (PGC) increased antlerless harvests to reduce deer density to meet deer management goals. Survival rates varied by month and age but not between study areas or among years after implementation of APRs. Monthly survival rates for subadults ranged from 0.64 to 0.97 during hunting seasons and 0.95 to 0.99 during the non-hunting period. Annual survival of subadults was 0.46 (95% CI = 0.41–0.52). Adult monthly survival rates ranged from 0.36 to 0.95 during hunting seasons and we had no mortalities during the non-hunting period. Annual survival of adults was 0.28 (95% CI = 0.22–0.35). Antler point restrictions successfully reduced harvest rate for subadults to 0.31 (95% CI = 0.23–0.38), and approximately 92% of these deer survived to the following hunting season. Vehicle collisions were the greatest source of mortality outside the hunting season for subadults and adults. Also, we observed decreased harvest rates for adults (0.59, 95% CI = 0.40–0.72), although nearly all were legal for harvest. Of radio-collared subadults, 6–11% were harvested with sub-legal antlers, indicating hunters generally complied with APRs. Overall, antlered harvest declined statewide and in our study areas, in part because of APRs but also because of increased antlerless harvests that reduced the statewide population from 1.49 million deer in 2000 to 1.14 million deer in 2005. However, between 2000 and 2005, harvest of adult males increased by 976 (112%) in Armstrong County, decreased by 29 (−3%) in Centre County, and increased by 14,285 (29%) statewide because more males survived to the 3- and 4-year-old age classes. Proportion of hunters from the random sample surveys who supported statewide APRs varied among years between 0.61 (95% CI = 0.59–0.64) and 0.70 (95% CI = 0.66–0.73). The proportion of hunters from the longitudinal panel who supported APRs did not increase as hunters gained experience under the new regulations; 0.23 were more supportive, 0.29 were less supportive, and 0.48 were unchanged in their level of agreement after 3 years. Although >50% of hunters supported APRs throughout the study, support for the PGC's deer management program declined; 41% of the longitudinal panel of hunters rated the deer management program lower after 3 years and 21% rated it higher. We considered APRs biologically successful because of decreased subadult harvest rates and increased harvest of adult males with larger antlers. Likewise, because the majority of hunters supported APRs throughout the study, we considered APRs socially successful. However, we predicted APRs would become increasingly popular after hunters experienced biological results of APRs, but there was little change in support. We believe hunters formed an initial impression of the effects of APRs, and additional experience and information failed to change their opinion. Furthermore, the concurrent reduction in overall deer densities to accommodate more males in the population and to meet agency deer population goals likely further reduced support for APRs. We found APRs as implemented in Pennsylvania to be enforceable, adhered to by hunters, and successful in recruiting more antlered males to older age classes. To facilitate social acceptance of these regulation changes, we found that obtaining support before the changes were implemented may have been important because most hunters did not change their opinions about APRs after 3 years of experience with the new regulations.

Pennsylvania

Evaluation of maternal penning to improve calf survival in the Chisana Caribou Herd

Predation is a major limiting factor for most small sedentary caribou ( Rangifer tarandus ) populations, particularly those that are threatened or endangered across the southern extent of the species’ range. Thus, reducing predation impacts is often a management goal for improving the status of small caribou populations, and lethal predator removal is the primary approach that has been applied. Given that predator control programs are often contentious, other management options that can garner broader public acceptance need to be considered. Substantial calf losses to predation in the few weeks following birth are common for these small caribou populations. Therefore, we employed a novel experimental approach of maternal penning with the goal of reducing early calf mortality in the Chisana Caribou Herd, a declining population in southwest Yukon and adjacent Alaska thought to number around 300 individuals. Maternal penning entailed temporarily holding pregnant females on their native range in a large pen secure from predators from late March through the initial weeks of calf rearing to mid‐June. During 2003–2006, we conducted 4 annual penning trials with 17–50 pregnant females each year ( n = 146 total), assessed survival of calves born in the pens, and evaluated survival and nutritional effects of penning for females that were held. We also investigated the herd's population dynamics during 2003–2008 to determine effects of maternal penning on calf recruitment and population growth. In addition to information gained during maternal penning, we determined natality and survival patterns via radiotelemetry, conducted autumn age‐sex composition surveys each year, and censused the population in mid‐October 2003, 2005, and 2007. Based on our penning trials and demographic investigations, we used simulation models to evaluate the effects of maternal penning relative to a population's inherent growth rate (finite rate of increase [λ] without maternal penning) and penning effort (proportion of calves born in penning) to provide perspective on utility of this approach for improving the status of small imperiled caribou populations. Pregnant females held in maternal penning tolerated captivity well in that they exhibited positive nutritional responses to ad libitum feed we provided and higher survival than free‐ranging females (0.993 and 0.951 for penned and free‐ranging females, respectively). Survival of pen calves from birth to mid‐June was substantially higher than that of free‐ranging calves ( = 0.950 and 0.376, respectively). This initial period accounted for 76% of the annual calf mortality in the free‐ranging population. Pen‐born calves maintained their survival advantage over wild‐born calves to the end of their first year ( = 0.575 and 0.192, respectively) during years penning occurred. Females in the Chisana Herd were highly productive with 57% producing their first offspring at 2 years of age, and annual natality rates averaging 0.842 calves/female ≥2 years old. Age‐specific natality rates exceeded 0.900 for 4–9‐year‐olds, then exhibited senescent decline to 0.467 by 19 years old. Annual survival of free‐ranging adult females and calves averaged 0.892 and 0.184, respectively, over all study years; both were reduced during 2004 because of poor winter survival. We noted reduced nutritional condition of caribou late that winter in that females we captured were lighter than in other years and produced lighter calves. We suspect that the reduced survival during winter 2004 and the observed nutritional characteristics resulted from adverse snow conditions in combination with effects of the extreme drought experienced the previous summer. Age‐specific survival of adult females was ≥0.900 through 10 years of age, then declined with age. The Chisana Herd numbered 720 caribou in mid‐October 2003, or more than twice that estimated prior to initiating maternal penning, and increased to 766 caribou by mid‐October 2007. We calculated that penning added 54.2 yearling recruits, or 40% of calves released from penning. Based on the maternal penning results and the population's vital rates, we determined that the herd would have been stable during 2003–2007 at about 713 caribou without maternal penning; thus, the increase in herd size we observed resulted from maternal penning and was equivalent to the estimate of additional yearling recruits. The improvement in the population trend invoked by maternal penning was limited by the larger than expected population size and resulting low penning effort ( = 11% of calves born in pen). Our simulations corroborated that maternal penning increased population size by the number of additional recruits provided, even at low penning effort, for inherently stable populations. As the inherent rate of increase dropped below λ = 1.000, more of the additional recruits from penning were needed to offset the downward population inertia, thus requiring increased penning effort to reach stability. For populations declining at λ < 0.890, stability could not be achieved with 100% penning effort given the vital rates in our models. Maternal penning in its limited application to date has proven to be broadly popular as a nonlethal management action aimed at reducing initial calf mortality from predation in small caribou populations. However, based on the Chisana program and 3 subsequent efforts elsewhere, improvement in population trends have been modest at best and come at a high financial cost. Given the necessity of maximizing penning effort, maternal penning may have a role in addressing conservation challenges for some small caribou populations that are stable or slowly declining, but its application should be primarily driven by objective assessment of the likelihood of improving population trends rather than popularity relative to other management options.

Wrangell‐St. Elias National Park and Preserve, Klu