White-tailed deer detection rates increase when coyotes are present.

Predator species can indirectly affect prey species through the cost of anti-predator behavior responses, which may involve shifts in occupancy, space use, or movement. Quantifying the various strategies implemented by prey species to avoid adverse interactions with predators can lead to a better understanding of potential population-level repercussions. Therefore, the purpose of this study was to examine predator-prey interactions by quantifying the effect of predator species presence on detection rates of prey species, using coyotes (Canis latrans) and white-tailed deer (Odocoileus virginianus) in Central Appalachian forests of the eastern United States as a model predator-prey system. To test two competing hypotheses related to interspecific interactions, we modeled species detections from 319 camera traps with a two-species occupancy model that incorporated a continuous-time detection process. We found that white-tailed deer occupancy was independent of coyote occupancy, but white-tailed deer were more frequently detectable and had greater detection intensity at sites where coyotes were present, regardless of vegetation-related covariates. In addition, white-tailed deer detection rates at sites with coyotes were highest when presumed forage availability was relatively low. These findings suggest that white-tailed deer may be exhibiting an active avoidance behavioral response to predators by increasing movement rates when coyotes are present in an area, perhaps due to reactive evasive maneuvers and/or proactive attempts to reduce adverse encounters with them. Concurrently, coyotes could be occupying sites with higher white-tailed deer densities. Because white-tailed deer did not exhibit significant shifts in daily activity patterns based on coyote occupancy, we further suggest that white-tailed deer in our study system generally do not use temporal partitioning as their primary strategy for avoiding encounters with coyotes. Overall, our study implements a recently developed analytical approach for modeling multi-species occupancy from camera traps and provides novel ecological insight into the complex relationships between predator and prey species.

Combining camera trap surveys and IUCN range maps to improve knowledge of species distributions.

Reliable maps of species distributions are fundamental for biodiversity research and conservation. The International Union for Conservation of Nature (IUCN) range maps are widely recognized as authoritative representations of species' geographic limits, yet they might not always align with actual occurrence data. In recent area of habitat (AOH) maps, areas that are not habitat have been removed from IUCN ranges to reduce commission errors, but their concordance with actual species occurrence also remains untested. We tested concordance between occurrences recorded in camera trap surveys and predicted occurrences from the IUCN and AOH maps for 510 medium- to large-bodied mammalian species in 80 camera trap sampling areas. Across all areas, cameras detected only 39% of species expected to occur based on IUCN ranges and AOH maps; 85% of the IUCN only mismatches occurred within 200 km of range edges. Only 4% of species occurrences were detected by cameras outside IUCN ranges. The probability of mismatches between cameras and the IUCN range was significantly higher for smaller-bodied mammals and habitat specialists in the Neotropics and Indomalaya and in areas with shorter canopy forests. Our findings suggest that range and AOH maps rarely underrepresent areas where species occur, but they may more often overrepresent ranges by including areas where a species may be absent, particularly at range edges. We suggest that combining range maps with data from ground-based biodiversity sensors, such as camera traps, provides a richer knowledge base for conservation mapping and planning.

Combinación de censos con fototrampas y mapas de extensión de la UICN para incrementar el conocimiento sobre la distribución de las especies Resumen Los mapas confiables de la distribución de las especies son fundamentales para la investigación y conservación de la biodiversidad. Los mapas de distribución de la Unión Internacional para la Conservación de la Naturaleza (UICN) están reconocidos como representaciones de autoridad de los límites geográficos de las especies, aunque no siempre se alinean con los datos actuales de su presencia. En los mapas recientes de área de hábitat (ADH), las áreas que no son hábitat han sido eliminadas de la distribución de la UICN para reducir los errores de comisión, pero su concordancia con la presencia actual de las especies tampoco ha sido analizada. Analizamos la concordancia entre la presencia registrada por los censos de fototrampas y pronosticamos la presencia a partir de los mapas de la UICN y de ADH de 510 especies de mamíferos de talla mediana a grande en 80 áreas de muestreo de fototrampas. Las cámaras detectaron sólo el 39% de las especies esperadas con base en la distribución de la UICN y los mapas de ADH en todas las áreas; el 85% de las disparidades con la UICN ocurrieron dentro de los 200 km a partir del borde de la distribución. Sólo el 4% de la presencia de las especies fue detectada por las cámaras ubicadas fuera de la distribución de la UICN. La probabilidad de disparidad entre las cámaras y la UICN fue significativamente mayor para los mamíferos de talla pequeña y para los especialistas de hábitat en las regiones Neotropical e Indomalaya y en áreas con doseles forestales más bajos. Nuestros hallazgos sugieren que los mapas de distribución y ADH pocas veces subrepresentan las áreas con presencia de las especies, pero con frecuencia pueden sobrerrepresentar la distribución al incluir áreas en donde las especies pueden estar ausentes, en particular los bordes de la distribución. Sugerimos que la combinación de los mapas de distribución con los sensores de biodiversidad en tierra, como las fototrampas, proporciona una base más rica de conocimiento para el mapeo y la planeación de la conservación.

Habitat management for stopover and breeding songbird communities along rights-of-way in forest-dominated landscapes.

The proliferation of energy rights-of-way (pipelines and powerlines; ROWs) in the central Appalachian region has prompted wildlife management agencies to consider ways to use these features to manage and conserve at-risk songbird species. However, little empirical evidence exists regarding best management strategies to enhance habitat surrounding ROWs for the songbird community during stopover or breeding periods. We used a before-after-control-impact design to study cut-back border (linear tree cuttings along abrupt forest edges) harvest width (15, 30, and 45 m wide into the forest), and harvest intensity (14 and 4.5 m2 /ha basal area retention) prescriptions along ROWs and assessed their effects on mature forest and young forest songbird species and avian guilds (forest gap habitat, forest interior habitat, young forest habitat, and species of regional conservation priority) up to 2 years after treatment throughout West Virginia. Species richness during the spring stopover period initially decreased at the 1-year post-treatment period but returned to pre-treatment levels by 2 years post-treatment. Breeding season responses to cut-back border treatments varied across harvest width, harvest intensity, and time, but all responses of focal species abundance and guild richness were neutral or positive. Cut-back border harvest intensity had a stronger influence (i.e., more positive responses) than harvest width on breeding focal species abundances and guild richness. For harvest intensity, the more intense, 4.5 m2 /ha retention treatment had a stronger influence (i.e., more positive responses) than the less intense, 14 m2 /ha retention treatment. For harvest width, the narrowest treatment (15-m wide) had the strongest influence (i.e., more positive responses) of all width treatments, followed by the widest (45-m wide treatment) with the least influence from the 30-m wide treatment. Abundances and richness increased from pre-treatment to 2 years post-treatment across all species and guilds that exhibited a response. These results suggest that cut-back borders increase breeding season habitat suitability along ROWs for the mature forest and young forest songbird community as well as for species of regional conservation priority in the short-term. These findings can aid the development of management guidelines for the forest songbird community along abrupt forest edges of man-made habitat features in forest-dominated landscapes.

A penalized likelihood for multispecies occupancy models improves predictions of species interactions.

Multispecies occupancy models estimate dependence among multiple species of interest from patterns of co-occurrence, but problems associated with separation and boundary estimates can lead to unreasonably large estimates of parameters and associated standard errors when species are rarely observed at the same site or when data are sparse. In this paper, we overcome these issues by implementing a penalized likelihood, which introduces a small bias in parameter estimates in exchange for a potentially large reduction in variance. We compare parameter estimates obtained from both penalized and unpenalized multispecies occupancy models fit to simulated data that exhibit various degrees of separation and to a real-word data set of bird surveys with little apparent overlap between potentially interacting species. Our simulation results demonstrate that penalized multispecies occupancy models did not exhibit boundary estimates and produced lower bias, lower mean squared error, and improved inference relative to unpenalized models. When applied to real-world data, our penalized multispecies occupancy model constrained boundary estimates and allowed for meaningful inference related to the interactions of two species of conservation concern. To facilitate the use of our penalized multispecies occupancy model, the techniques demonstrated in this paper have been integrated into the unmarked package in R programing language.

Factors influencing productivity of eastern wild turkeys in northeastern South Dakota.

Population growth is highly sensitive to changes in reproductive rates for many avian species. Understanding how reproductive rates are related to environmental conditions can give managers insight into factors contributing to population change. Harvest trends of eastern wild turkey in northeastern South Dakota suggest a decline in abundance. We investigated factors influencing reproductive success of this important game bird to identify potential factors contributing to the decline. We monitored nesting rate, nest survival, renesting rate, clutch size, hatchability, and poult survival of 116 eastern wild turkey hens using VHF radio transmitters during the springs and summers of 2017 and 2018. Heavier hens were more likely to attempt to nest than lighter hens, and adult hens were more likely to renest than yearling hens. Nest survival probability was lowest in agricultural fields relative to all other cover types and positively related to horizontal visual obstruction and distance to the nearest road. Daily nest survival probability demonstrated an interaction between temperature and precipitation, such that nest survival probability was lower on warm, wet days, but lowest on dry days. Egg predation was the leading cause of nest failure, followed by haying of the nest bowl and death of the incubating hen. Poults reared by adult hens had a greater probability of survival than poults reared by yearling hens. Our estimate of survival probability of poults raised by yearling hens was low relative to other studies, which may be contributing to the apparent regional population decline. However, there is little managers can do to influence poult survival in yearling hens. Alternatively, we found nest survival probability was lowest for nests initiated in agricultural fields. Wildlife-friendly harvesting practices such as delayed haying or installation of flushing bars could help increase productivity of eastern wild turkey in northeastern South Dakota.

A comparison of wetland characteristics between Agricultural Conservation Easement Program and public lands wetlands in West Virginia, USA.

In West Virginia, USA, there are 24 conservation easement program wetlands enrolled in the Agricultural Conservation Easement Program (ACEP). These wetlands are located on private agricultural land and are passively managed. Due to their location within fragmented agricultural areas, wetlands enrolled in ACEP in West Virginia have the potential to add wetland ecosystem services in areas that are lacking these features. We evaluated ACEP wetlands compared to reference wetlands on public land in West Virginia by using surrounding land cover, vegetative cover, and wetland features and stressors such as the presence or absence of erosion, upland inclusion, algal mats, and evidence of impacts from the surrounding landscape as surrogate measurements of wetland function on 13 ACEP wetlands and 10 reference wetlands. ACEP wetlands had higher percentages of tree coverage and a higher proportion of agricultural land in the areas immediately surrounding the wetland. Reference wetlands had higher percent coverage of emergent vegetation and had a higher proportion of forest in the immediate landscape. Our findings suggest that ACEP wetlands provide valuable early successional and forested wetland cover in a state that is largely forested. Because of this, it is important to maintain and even expand ACEP in West Virginia to continue providing a valuable source of early successional wetland habitat.

Effects of forest management on vertebrates: synthesizing two decades of data from hardwood forests in Missouri, USA.

Modern forest management seeks to balance multiple social, economic, and ecological goals. Different management approaches create different types of disturbances in a forest ecosystem and thus also differ in their impacts on plants, animals, and insects. Understanding these impacts is important for conservation of forest ecosystem function, but challenging due to the large spatial and temporal scale over which management occurs. Most past research has focused on relatively small areas, short time scales, and/or a small number of species. To address this, we examined the effects of two common silvicultural systems (even and uneven aged) on abundance and richness of three vertebrate taxa (birds, small mammals, and herpetofauna) over a two-decade period in a temperate hardwood forest in Missouri, USA. The two systems removed a similar amount of biomass overall, but differed in the intensity, number, and configuration of harvests applied. We found that vertebrate population responses varied by taxa, occurred at multiple spatial scales, and were concentrated in the period following the first harvest entry. Birds generally had the largest changes in relative abundance, both positive and negative, following management. Small mammals and reptiles had smaller, but generally positive, responses; amphibians were mixed. Bird species tended to respond in the same way to both silvicultural systems, while small mammals and herpetofauna did not respond consistently. Thus, for birds, the total amount of harvest disturbance across the landscape drives population responses, while for others the size and configuration of individual harvests is likely more important. Synthesizing results across the vertebrate community at large spatial and temporal scales allows managers to better understand trade-offs when making decisions that will affect wildlife in contrasting ways.

Nutritional condition and physiological stress levels of elk in the Black Hills, South Dakota.

Percent of body fat and physiological stress are important correlates to wildlife demographics. We studied winter percent of body fat and physiological stress levels for a declining elk (Cervus canadensis nelsoni) population in South Dakota, 2011-2013. We obtained percent of winter body fat, pregnancy status, lactation status, and physiological stress data from 58 adult females (2+ years old). We compared physiological stress level data from 2011 with data collected from this same herd when elk densities were much higher (1995-1997). Our objectives were to determine percent of body fat during winter, examine if winter body fat was correlated with pregnancy and lactation status, and quantify and compare physiological stress hormone values from elk in the mid-1990s. Probability of being pregnant increased with higher winter nutritional condition, or percent of body fat; whereas females with a higher probability of previously lactating were lower in winter body fat. Mean fecal glucocorticoid metabolite (FGM) levels in 2011 (mean = 47.78 ng/g, SE = 2.37) were higher during summer compared to data collected in 1995-1997 (mean = 34.21 ng/g, SE = 3.71); however, mean FGM levels during winter did not differ between the two time periods. Although summer levels of FGM have significantly increased since the mid-1990s, we caution against any interpretation of increased FGM levels on elk fitness, as it may not infer biological significance. Mean winter percent of body fat of elk was lower when compared to other populations in the west but this difference does not appear to be limiting vital rates and population growth for this elk herd. We recommend future research focus on summer/autumn data collection to provide a more comprehensive understanding of percent of body fat for elk in our region.

Influence of the Agricultural Conservation Easement Program wetland practices on winter occupancy of Passerellidae sparrows and avian species richness.

Wetlands enrolled in the Agricultural Conservation Easement Program (ACEP) are established as a means of restoring wetland ecosystems and wildlife habitat on private, agricultural land. In West Virginia, USA, ACEP wetlands have never been evaluated to determine how they function as wildlife habitat in comparison to other available wetland habitat in the state. We measured the wintering occupancy of Passerellidae species and apparent avian species richness on ACEP wetlands and a set of reference wetlands located on public land in West Virginia to evaluate if ACEP wetlands are being used similarly by avian species to other available wetland habitat in the state. Apparent avian species richness and the occupancy probability of four Passerellidae species-song sparrows (Melospiza melodia), dark-eyed juncos (Junco hyemalis), swamp sparrows (Melospiza georgiana), and white-throated sparrows (Zonotrichia albicollis)-did not differ between ACEP and reference sites. In addition to other vegetative and habitat associations for each species, dark-eyed junco occupancy was negatively correlated with wetland size while swamp sparrow occupancy and apparent avian species richness were positively associated with wetland size. These results indicate that ACEP wetlands are providing winter avian habitat as well as another source of wetland habitat in the state. Maintaining and expanding ACEP wetlands in West Virginia would continue to provide wetland systems in areas that are otherwise lacking these habitats.

Mammal communities are larger and more diverse in moderately developed areas.

Developed areas are thought to have low species diversity, low animal abundance, few native predators, and thus low resilience and ecological function. Working with citizen scientist volunteers to survey mammals at 1427 sites across two development gradients (wild-rural-exurban-suburban-urban) and four plot types (large forests, small forest fragments, open areas and residential yards) in the eastern US, we show that developed areas actually had significantly higher or statistically similar mammalian occupancy, relative abundance, richness and diversity compared to wild areas. However, although some animals can thrive in suburbia, conservation of wild areas and preservation of green space within cities are needed to protect sensitive species and to give all species the chance to adapt and persist in the Anthropocene.

Correction: Seeing the Forest through the Trees: Considering Roost-Site Selection at Multiple Spatial Scales.

[This corrects the article DOI: 10.1371/journal.pone.0150011.].

Simulated data and code for analysis of herpetofauna response to forest management in the Missouri Ozarks.

We present predictor variables and R and Stan code for simulating and analyzing counts of Missouri Ozark herpetofauna in response to three forest management strategies. Our code performs four primary purposes: import predictor variables from spreadsheets; simulate synthetic response variables based on imported predictor variables and user-supplied values for data-generating parameters; format synthetic data for export to Stan; and analyze synthetic data.

Seeing the Forest through the Trees: Considering Roost-Site Selection at Multiple Spatial Scales.

Conservation of bat species is one of the most daunting wildlife conservation challenges in North America, requiring detailed knowledge about their ecology to guide conservation efforts. Outside of the hibernating season, bats in temperate forest environments spend their diurnal time in day-roosts. In addition to simple shelter, summer roost availability is as critical as maternity sites and maintaining social group contact. To date, a major focus of bat conservation has concentrated on conserving individual roost sites, with comparatively less focus on the role that broader habitat conditions contribute towards roost-site selection. We evaluated roost-site selection by a northern population of federally-endangered Indiana bats (Myotis sodalis) at Fort Drum Military Installation in New York, USA at three different spatial scales: landscape, forest stand, and individual tree level. During 2007-2011, we radiotracked 33 Indiana bats (10 males, 23 females) and located 348 roosting events in 116 unique roost trees. At the landscape scale, bat roost-site selection was positively associated with northern mixed forest, increased slope, and greater distance from human development. At the stand scale, we observed subtle differences in roost site selection based on sex and season, but roost selection was generally positively associated with larger stands with a higher basal area, larger tree diameter, and a greater sugar maple (Acer saccharum) component. We observed no distinct trends of roosts being near high-quality foraging areas of water and forest edges. At the tree scale, roosts were typically in American elm (Ulmus americana) or sugar maple of large diameter (>30 cm) of moderate decay with loose bark. Collectively, our results highlight the importance of considering day roost needs simultaneously across multiple spatial scales. Size and decay class of individual roosts are key ecological attributes for the Indiana bat, however, larger-scale stand structural components that are products of past and current land use interacting with environmental aspects such as landform also are important factors influencing roost-tree selection patterns.

A two-species occupancy model accommodating simultaneous spatial and interspecific dependence.

Occupancy models are popular for estimating the probability a site is occupied by a species of interest when detection is imperfect. Occupancy models have been extended to account for interacting species and spatial dependence but cannot presently allow both factors to act simultaneously. We propose a two-species occupancy model that accommodates both interspecific and spatial dependence. We use a point-referenced multivariate hierarchical spatial model to account for both spatial and interspecific dependence. We model spatial random effects with predictive process models and use probit regression to improve efficiency of posterior sampling. We model occupancy probabilities of red fox (Vulpes vulpes) and coyote (Canis latrans) with camera trap data collected from six mid-Atlantic states in the eastern United States. We fit four models comprising a fully factorial combination of spatial and interspecific dependence to two-thirds of camera trapping sites and validated models with the remaining data. Red fox and coyotes each exhibited spatial dependence at distances > 0.8 and 0.4 km, respectively, and exhibited geographic variation in interspecific dependence. Consequently, predictions from the model assuming simultaneous spatial and interspecific dependence best matched test data observations. This application highlights the utility of simultaneously accounting for spatial and interspecific dependence.

The role of wildfire, prescribed fire, and mountain pine beetle infestations on the population dynamics of black-backed woodpeckers in the black hills, South Dakota.

Wildfire and mountain pine beetle infestations are naturally occurring disturbances in western North American forests. Black-backed woodpeckers (Picoides arcticus) are emblematic of the role these disturbances play in creating wildlife habitat, since they are strongly associated with recently-killed forests. However, management practices aimed at reducing the economic impact of natural disturbances can result in habitat loss for this species. Although black-backed woodpeckers occupy habitats created by wildfire, prescribed fire, and mountain pine beetle infestations, the relative value of these habitats remains unknown. We studied habitat-specific adult and juvenile survival probabilities and reproductive rates between April 2008 and August 2012 in the Black Hills, South Dakota. We estimated habitat-specific adult and juvenile survival probability with Bayesian multi-state models and habitat-specific reproductive success with Bayesian nest survival models. We calculated asymptotic population growth rates from estimated demographic rates with matrix projection models. Adult and juvenile survival and nest success were highest in habitat created by summer wildfire, intermediate in MPB infestations, and lowest in habitat created by fall prescribed fire. Mean posterior distributions of population growth rates indicated growing populations in habitat created by summer wildfire and declining populations in fall prescribed fire and mountain pine beetle infestations. Our finding that population growth rates were positive only in habitat created by summer wildfire underscores the need to maintain early post-wildfire habitat across the landscape. The lower growth rates in fall prescribed fire and MPB infestations may be attributed to differences in predator communities and food resources relative to summer wildfire.

A re-evaluation of a case-control model with contaminated controls for resource selection studies.

1. A common sampling design in resource selection studies involves measuring resource attributes at sample units used by an animal and at sample units considered available for use. Few models can estimate the absolute probability of using a sample unit from such data, but such approaches are generally preferred over statistical methods that estimate a relative probability of use. 2. The case-control model that allows for contaminated controls, proposed by Lancaster & Imbens (1996) and Lele (2009), can estimate the absolute probability of using a sample unit from use-availability data. However, numerous misconceptions have likely prevented the widespread application of this model to resource selection studies. We address common misconceptions regarding the case-control model with contaminated controls and demonstrate its ability to estimate the absolute probability of use, prevalence and parameters associated with categorical covariates from use-availability data. 3. We fit the case-control model with contaminated controls to simulated data with varying prevalence (defined as the average probability of use across all sample units) and sample sizes (n1 = 500 used and na = 500 available samples; n1 = 1000 used and na = 1000 available samples). We then applied this model to estimate the probability Ozark hellbenders (Cryptobranchus alleganiensis bishopi) would use a location within a stream as a function of covariates. 4. The case-control model with contaminated controls provided unbiased estimates of all parameters at N = 2000 sample size simulation scenarios, particularly at low prevalence. However, this model produced increasingly variable maximum likelihood estimates of parameters as prevalence increased, particularly at N = 1000 sample size scenarios. We thus recommend at least 500-1000 used samples when fitting the case-control model with contaminated controls to use-availability data. Our application to hellbender data revealed selection for locations with coarse substrate that are close to potential sources of cover. 5. This study unites a disparate literature, addresses and clarifies many commonly held misconceptions and demonstrates that the case-control model with contaminated controls is a viable alternative for estimating the absolute probability of use from use-availability data.

Predicting species distributions from samples collected along roadsides.

Predictive models of species distributions are typically developed with data collected along roads. Roadside sampling may provide a biased (nonrandom) sample; however, it is currently unknown whether roadside sampling limits the accuracy of predictions generated by species distribution models. We tested whether roadside sampling affects the accuracy of predictions generated by species distribution models by using a prospective sampling strategy designed specifically to address this issue. We built models from roadside data and validated model predictions at paired locations on unpaved roads and 200 m away from roads (off road), spatially and temporally independent from the data used for model building. We predicted species distributions of 15 bird species on the basis of point-count data from a landbird monitoring program in Montana and Idaho (U.S.A.). We used hierarchical occupancy models to account for imperfect detection. We expected predictions of species distributions derived from roadside-sampling data would be less accurate when validated with data from off-road sampling than when it was validated with data from roadside sampling and that model accuracy would be differentially affected by whether species were generalists, associated with edges, or associated with interior forest. Model performance measures (kappa, area under the curve of a receiver operating characteristic plot, and true skill statistic) did not differ between model predictions of roadside and off-road distributions of species. Furthermore, performance measures did not differ among edge, generalist, and interior species, despite a difference in vegetation structure along roadsides and off road and that 2 of the 15 species were more likely to occur along roadsides. If the range of environmental gradients is surveyed in roadside-sampling efforts, our results suggest that surveys along unpaved roads can be a valuable, unbiased source of information for species distribution models.