Publications by Colleges and Departments (MSU - Bozeman)
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Item Responses of American black bears to spring resources(Wiley, 2021-11) Bowersock, Nathaniel R.; Litt, Andrea R.; Merkle, Jerod A.; Gunther, Kerry A.; van Manen, Frank T.In temperate regions of the world, food resources are seasonally limited, which causes some wildlife species to seek out nutrient-rich resources to better meet their caloric needs. Animals that utilize high-quality resources may reap fitness benefits as they prepare for mating, migration, or hibernation. American black bears (Ursus americanus) are omnivores that consume both plant and animal food resources to meet macronutrient needs. Black bears capitalize on high-quality food resources, such as soft mast in summer and hard mast during autumn, but we know less about the importance of resource quality during spring. Therefore, we sought to understand the relationship between the spatiotemporal variation in the availability of food and resource selection of black bears during spring. We also aimed to infer potential changes in foraging tactics, from opportunistic foraging to more active selection. Although black bears are described as opportunistic omnivores, we hypothesized they select areas with high-quality forage when available. We instrumented 7 black bears with GPS collars in 2017 and 2018 and estimated fine-scale resource selection with integrated step-selection functions. We found evidence that black bear movements were influenced by forage quality of vegetative food resources. However, we failed to find evidence that black bears actively alter their movements to take advantage of seasonal neonate elk. Although black bears represent a substantial cause of mortality for neonate elk, we found that black bears likely feed on neonates encountered opportunistically while traveling between patches of high-quality forage. Few studies have shown evidence of an omnivorous species capitalizing on spatiotemporal variation in forage quality, yet our data suggest this may be an important strategy for species with diverse diets, particularly where resources are seasonally limited.Item Rub tree use and selection by American black bears and grizzly bears in northern Yellowstone National Park(Ursus, 2022-06) Bowersock, Nathaniel R.; Okada, Hitomi; Litt, Andrea R.; Gunther, Kerry A.; van Manen, Frank T.Several of the world's bear species exhibit tree-rubbing behavior, which is thought to be a form of scent-marking communication. Many aspects of this behavior remain unexplored, including differences in rub tree selection between sympatric bear species. We compiled rub tree data collected on Yellowstone National Park's Northern Range (USA) and compared rub tree selection of sympatric American black bears (Ursus americanus) and grizzly bears (U. arctos) at local and landscape scales. During 2017 and 2018, we identified 217 rub trees and detected black bears at 117 rub trees and grizzly bears at 18 rub trees, based on genetic analysis of collected hair samples. Rub trees generally were located in areas with gentle slopes and close to existing animal trails. Trees selected by black bears were typically in forested areas, whereas trees selected by grizzly bears were in forested and more open areas. Use of rub trees varied seasonally and between sexes for black bears, but seasonal data were inconclusive for grizzly bears. Black bears showed preferences for certain tree species for rubbing, but we did not find evidence that rub tree selection by grizzly bears differed among tree species. Both bear species selected trees that lacked branches on the lower portions of tree trunks and the maximum rub height was consistent with the body length of the bear species that used the tree. Although the sample size for grizzly bears was small, identifying the species and sex of bears based on genetic analysis enhanced interpretation of rub tree use and selection by bears. Scent-marking by black bears and grizzly bears on similar rub objects in well-traversed areas likely serves to enhance communication within and between the 2 species.Item A functional ecology framework for understanding and predicting animal responses to plant invasion(Springer Science and Business Media LLC, 2022-05) Litt, Andrea R.; Pearson, Dean E.Plant invasions can alter food resources and habitat conditions that structure animal communities. These effects are negative for many native animals, but neutral or even positive for others. Understanding why we see this variation in responses is critical for mitigating invasion outcomes, yet we lack a synthetic framework to explain and potentially predict effects of invasive plants on native animals. We propose a trait-based framework for understanding how invasive plants affect native fauna, which draws on community assembly, niche, and trait theories to define the mechanisms by which invasive plants alter ecological conditions relevant to native animals. This approach moves beyond prior frameworks by explicitly accounting for the context dependency that defines most ecological interactions and invasion outcomes. Namely, by characterizing the plant community in terms of functional effect traits (e.g., seed size) relevant to consumers and quantifying those traits along a consumer resource axis, we can map the functional relationship between plant resources and animals. We can then delineate how plant invaders alter the plant community and associated resource axes to restructure consumer communities. We apply this framework to case studies of rodents, spiders, and birds to demonstrate the process and explore its utility. For example, we show that by focusing on how a nonnative grass altered seed sizes (relative to the native plant community), we can better understand declines in abundance of granivorous rodents and increases in opportunists. This approach can elucidate which native animals will be most likely affected by plant invasion, as well as how and why they might respond. Moreover, these mechanistic explanations provide working hypotheses for how invasive plants impact native animals more generally, with potential for predicting impacts of future invaders.Item Improving geographically extensive acoustic survey designs for modeling species occurrence with imperfect detection and misidentification(2018-06) Banner, Katharine M.; Irvine, Kathryn M.; Rodhouse, Thomas J.; Wright, Wilson J.; Rodriguez, Rogelio M.; Litt, Andrea R.Acoustic recording units (ARUs) enable geographically extensive surveys of sensitive and elusive species. However, a hidden cost of using ARU data for modeling species occupancy is that prohibitive amounts of human verification may be required to correct species identifications made from automated software. Bat acoustic studies exemplify this challenge because large volumes of echolocation calls could be recorded and automatically classified to species. The standard occupancy model requires aggregating verified recordings to construct confirmed detection/non-detection datasets. The multistep data processing workflow is not necessarily transparent nor consistent among studies. We share a workflow diagramming strategy that could provide coherency among practitioners. A false-positive occupancy model is explored that accounts for misclassification errors and enables potential reduction in the number of confirmed detections. Simulations informed by real data were used to evaluate how much confirmation effort could be reduced without sacrificing site occupancy and detection error estimator bias and precision. We found even under a 50% reduction in total confirmation effort, estimator properties were reasonable for our assumed survey design, species-specific parameter values, and desired precision. For transferability, a fully documented r package, OCacoustic, for implementing a false-positive occupancy model is provided. Practitioners can apply OCacoustic to optimize their own study design (required sample sizes, number of visits, and confirmation scenarios) for properly implementing a false-positive occupancy model with bat or other wildlife acoustic data. Additionally, our work highlights the importance of clearly defining research objectives and data processing strategies at the outset to align the study design with desired statistical inferences.Item Habitat characteristics of the hoary marmot: assessing distribution limitations in Montana(2018-10) Turnock, Benjamin Y.; Litt, Andrea R.; Vore, John M.; Hammond, C. A. M.Species that live in ecosystems with extremely different seasonal conditions must balance the constraints of each season to survive. Alpine species that do not migrate seasonally are especially adept at balancing the constraints created by short growing seasons and long, harsh winters. We investigated the habitat characteristics of hoary marmots in western Montana to provide a better understanding about habitat selection at the southern extent of this species' distribution. Hoary marmots are an alpine obligate of special concern in western Montana; given that climate change is impacting alpine ecosystems at a rapid rate, this species may be especially vulnerable at the southern edge of their range. We conducted occupancy surveys in three study areas along a latitudinal gradient in 2014 and 2015 to assess the importance of specific habitat characteristics to their presence on the landscape. Slope, aspect, and presence of shrubs were all important habitat characteristics. Marmots preferred shallow slopes and southern aspects, similar to findings from other studies on hoary marmots and other marmot species. Our results provide evidence that marmots may strike a balance between the environmental conditions they require during summer and winter. Shallow slopes typically accumulate deeper snow in winter that provide the best insulating snowpack. However, a preference for southern aspects allows for more snow-free areas in spring, providing a slightly longer growing season than northern aspects. Hoary marmots may be selecting areas with shrubs because shrubs can accumulate deeper snow and the additional insulation can increase subnivian temperatures. Other studies suggest that marmot survival is influenced by snowpack, indicating that marmot distribution may be more closely tied to winter conditions rather than summer conditions. This highlights the difficulty of working on marmots and other alpine obligates, as most studies occur only during the short growing season. Given the current and projected increases in temperature and reduction in snowpack in Montana, areas that provide the winter conditions hoary marmots require may become more limited. Effectively conserving, monitoring, and managing alpine obligates under an uncertain climate future will require a closer look at how winter conditions drive habitat selection and distributions on the landscape.Item Movement of Imperiled Chiricahua Leopard Frogs during Summer Monsoons(2018-12) Hinderer, Ross K.; Litt, Andrea R.; McCaffery, MagnusHabitat fragmentation and subsequent disruption of animal movement are responsible for extinctions in some species, including amphibians. Amphibians that can travel across fragmented landscapes may be at lower risk of extinction. Chiricahua Leopard Frogs are threatened because of fragmentation associated with habitat loss and degradation. We sought to identify cues leading frogs to leave perennial ponds and factors related to movement ability of Chiricahua Leopard Frogs in New Mexico during the summers of 2013 and 2014. Using pitfall traps, we captured frogs leaving ponds and radio-tracked 30 individuals to characterize overland movements. We checked traps and located frogs daily for up to eight weeks. We assessed factors related to the number of frogs leaving ponds using linear models and to distances moved by frogs using linear mixed models. The number of frogs caught was related to rainfall, but not water temperature, and more frogs were found outside ponds at an intermediate level of rainfall. Frogs who left ponds moved an average of 97 m/ day, but distances were highly variable among individuals. Sex and size did not explain differences in distances moved, after accounting for individual variation. One individual moved 1,658 m in a day and another 9,888 m over 36 days. These distances are farther than recorded previously for this species. Movement data are essential for developing recovery plans for threatened species, and our findings will inform planning by predicting the ability of populations to cope with the effects of habitat fragmentation and the ability of animals to colonize new habitats.Item Simulating cheatgrass (Bromus tectorum) invasion decreases access to food resources for small mammals in sagebrush steppe(2018-03) Bachen, Daniel A.; Litt, Andrea R.; Gower, Claire N.Invasions by nonnative plants can alter the abundance of native animals, yet we know little about the mechanisms driving these changes. Shifts in vegetation characteristics resulting from nonnative plants can alter availability of food resources, predation risk, and foraging efficiency (both the access to and ability to find food), each providing a potential mechanism for documented changes in animal communities and populations in invaded systems. Cheatgrass (Bromus tectorum) is a nonnative grass that invades sagebrush steppe, resulting in declines in some small mammal populations. We examined whether changes in structural characteristics associated with cheatgrass invasion could alter foraging by small mammals, providing a potential mechanism for documented population declines. We quantified differences in vegetation structure between native and cheatgrass-invaded sagebrush steppe, then experimentally added artificial structure in native areas to simulate these differences. We placed grain at foraging stations and measured the amount removed by small mammals nightly. Adding litter at depths approximating invasion by cheatgrass reduced the average amount of grain removed in 2 of 3 study areas, but increasing stem density did not. Based on this experiment, the deeper litter created by cheatgrass invasion may increase costs to small mammals by decreasing foraging efficiency and access to existing food resources, which may explain population-level declines in small mammals documented in other studies. By isolating and identifying which structural attributes of cheatgrass invasion are most problematic for small mammals, land managers may be able to design treatments to efficiently mitigate impacts and restore invaded ecosystems.Item A Detection Probability Model for Aerial Surveys of Mule Deer(2016-11) Zabransky, Cody J.; Hewitt, David G.; Deyoung, Randy W.; Gray, Shawn S.; Richardson, Calvin; Litt, Andrea R.; Deyoung, Charles A.Population estimates derived from aerial surveys of ungulates are biased by imperfect detection, where probability of sighting groups is influenced by variables specific to terrain features and vegetation communities. Therefore, methods for bias-correction must be validated for the region to which they will be applied. Our objectives were to quantify factors affecting detection probability of mule deer (Odocoileus hemionus) during helicopter surveys in Texas, USA, rangelands, and develop a detection probability model to reduce bias in deer population estimates. We placed global positioning system (GPS) collars on 215 deer on 6 sites representative of mule deer range in the southern Great Plains and the Chihuahuan Desert during 2008-2010. We collected data during aerial surveys in January-March and fit logistic regression models to predict detection probability of mule deer based on ecological and behavioral covariates. We evaluated the model using independent estimates of population size derived from a mark-resight procedure. Detection of mule deer was negatively related to distance from the transect, increasing brush cover, sunlight, and increasing terrain ruggedness (P< 0.01). Probability of detection in brush cover was greater if deer were active (P = 0.02). Population estimates corrected for visibility bias using our detection probability model or mark-resight models averaged 2.1 +/- 0.49 (SD; n = 50) and 2.2 +/- 0.62 times larger, respectively, than uncorrected counts. Estimates of population size derived from the detection probability model averaged 101 +/- 26% of mark-resight estimates. However, the detection probability model did not improve precision of population estimates, probably because of unmodeled variation in availability of deer during surveys. Our detection probability model is a simple and effective means to reduce bias in estimates of mule deer population size in southwestern rangelands. Availability bias may be a persistent issue for surveys of mule deer in the Southwest, and appears to be a primary influence of variance of population estimates.Item Performance of Juvenile Cutthroat Trout Translocated as Embryos from Five Populations into a Common Habitat(2016-07) Andrews, Tessa; Shepard, Bradley B.; Litt, Andrea R.; Kruse, Carter G.; Nelson, M. Lee; Clancey, Patrick; Zale, Alexander V.; Taper, Mark L.; Kalinowski, Steven T.The distributions of most native trout species in western North America have been severely reduced, and conservation of many of these species will require translocation into vacant habitats following removal of nonnative species. A critical question managers have is \Does it matter which donor sources are used for these translocations?\" We present a case study that addressed this question for a large native trout translocation project in Montana. We introduced embryos from five source populations of Westslope Cutthroat Trout Oncorhynchus clarkii lewisi to a large, fishless watershed in Montana following removal of nonnative fish with piscicides. Source populations providing embryos for translocations were three nearby (< 120 km) wild populations, the state of Montana\'s captive Westslope Cutthroat Trout hatchery conservation population (initiated 32 years ago using fish from wild populations located > 350 km from the translocation site), and a population in captivity for one generation comprised of individuals from the three wild populations used as single sources for this project, which were variably crossed (59% within populations and 41% between populations) to provide embryos. We used remote-site incubators at six different sites to introduce approximately 35,000 embryos from 400 genotyped parents. We later resampled and genotyped 1,450 of these individuals at age 1 and age 2. Juvenile survival for the more genetically diverse Montana Westslope Cutthroat Trout conservation population was twice as high as for other source populations, even though these other source populations were geographically closer to the translocation site than populations used to make the Montana Westslope Cutthroat Trout conservation population. Body weight for progeny from the two captive populations was higher than for progeny from wild source populations, and some differences were observed in body condition among source populations. Continued monitoring over several generations will be necessary to determine the eventual contributions of each source population and the relevance of these initial findings."Item Larval long-toed salamanders incur nonconsumptive effects in the presence of nonnative trout(2016-05) Kenison, Erin K.; Litt, Andrea R.; Pilliod, David S.; McMahon, Thomas E.Predators can influence prey directly through consumption or indirectly through nonconsumptive effects (NCEs) by altering prey behavior, morphology, and life history. We investigated whether predator-avoidance behaviors by larval long-toed salamanders (Ambystoma macrodactylum) in lakes with nonnative trout result in NCEs on morphology and development. Field studies in lakes with and without trout were corroborated by experimental enclosures, where prey were exposed only to visual and chemical cues of predators. We found that salamanders in lakes with trout were consistently smaller than in lakes without trout: 38% lower weight, 24% shorter body length, and 29% shorter tail length. Similarly, salamanders in protective enclosures grew 2.9 times slower when exposed to visual and olfactory trout cues than when no trout cues were present. Salamanders in trout-free lakes and enclosures were 22.7 times and 1.48 times, respectively, more likely to metamorphose during the summer season than those exposed to trout in lakes and/or their cues. Observed changes in larval growth rate and development likely resulted from a facultative response to predator-avoidance behavior and demonstrate NCEs occurred even when predation risk was only perceived. Reduced body size and growth, as well as delayed metamorphosis, could have ecological consequences for salamander populations existing with fish if those effects carry-over into lower recruitment, survival, and fecundity.