Olfactory and visual plant cues as drivers of selective herbivory

Food quality is an important consideration in the foraging strategy of all animals, including herbivores. Those that can detect and assess the nutritional value of plants from afar, using senses such as smell and sight, can forage more efficiently than those that must assess food quality by taste alone. Selective foraging not only affects herbivore fitness but can influence the structure and composition of plant communities, yet little is known about how olfactory and visual cues help herbivores to find preferred plants. We tested the ability of a free‐ranging, generalist mammalian browser, the swamp wallaby Wallabia bicolor, to use olfactory and visual plant cues to find and/or browse differentially on Eucalyptus pilularis seedlings grown under different nutrient conditions. Low‐nutrient seedlings differed from high‐nutrient seedlings, having lighter coloured leaves, red stems and lower biomass and nitrogen content. In the absence of visual cues, wallabies used odour to differentiate vials containing cut seedlings. They visited and investigated patches with high‐nutrient seedling odour most, followed by patches with low‐nutrient seedling odour, and patches with no added odour least. However, when visual and olfactory cues of seedlings were present, wallabies reversed their foraging response and were more likely to browse low‐ than high‐nutrient seedlings. This browsing difference, in turn, disappeared when long‐range visual cues were reduced by pinning seedlings horizontal to the ground. We suggest that visual cues overrode the effects of olfactory cues on browsing patterns of intact seedlings. Our study shows that herbivores can respond to odours of higher nutrient plants but in ecologically realistic scenarios they use a variety of visual and olfactory cues, with a context‐dependent outcome that is not always selection of high nutrient food. Our results demonstrate the importance of testing the sensory abilities of herbivores in realistic multi‐sensory settings to understand their function in selective foraging.     

Human activity and habitat type affect perceived predation risk in urban white‐footed mice (Peromyscus leucopus)

Predation risk is one of the largest costs associated with foraging in small mammals. Small mammals select microhabitat features such as tree and shrub canopy cover, woody debris and vegetative ground cover that can lower the risk of detection from predators and provide greater protection if discovered. Small mammals also increase foraging activity and decrease selection for cover when cloud cover increases and moon illumination is less. Often researchers assume small mammals in urban areas respond to these cues in the same manner as in natural areas, but these cues themselves are altered in urban zones. In this study, we investigated how Amur honeysuckle (Lonicera maackii) and coarse woody debris (CWD) affected giving‐up density (GUD) in white‐footed mice (Peromyscus leucopus). Each of three habitat treatments (open flood channel, the edge and interior of the honeysuckle patch) contained cover treatments with coarse woody debris present or absent. The six treatment combinations were compared to environmental variables (temperature, humidity and illumination) and habitat variables to test their effect on GUD. Peromyscus leucopus foraged to lower densities in areas with CWD present and also under the honeysuckle canopy, using this invasive shrub to decrease predation risk, potentially increasing survivability within this urban park. Increased human presence negatively affected foraging behavior across treatments. Human presence and light pollution significantly influenced P. leucopus, modifying their foraging behavior and demonstrating that both fine‐ and coarse‐scale urban factors can affect small mammals. Foraging increased as humidity increased, particularly under the honeysuckle canopy. Changes in illumination due to moonlight and cloud cover did not affect foraging behavior, suggesting urban light pollution may have altered behavioral responses to changes in light levels. Lonicera maackii seemed to facilitate foraging in P. leucopus, even though it adversely affects the plant community, suggesting that its impact may not be entirely negative.     

Spatial context influences patch residence time in foraging hierarchies

Understanding responses of organisms to spatial heterogeneity in resources has emerged as a fundamentally important challenge in contemporary ecology. We examined responses of foraging herbivores to multi-scale heterogeneity in plants. We asked the question, “Is the behavior observed at coarse scales in a patch hierarchy the collective outcome of fine scale behaviors or, alternatively, does the spatial context at coarse scales entrain fine scale behavior?” To address this question we created a nested, two-level patch hierarchy. We examined the effects of the spatial context surrounding a patch on the amount of time herbivores resided in the patch. We developed a set of competing models predicting residence time as a function of the mass of plants contained in a patch and the distance between patches and examined the strength of evidence in our observations for these models. Models that included patch mass and inter-patch distance as independent variables successfully predicted observed residence times (bears: r ²=0.67–0.76 and mule deer: r ²=0.33–0.55). Residence times of grizzly bears (Ursus arctos) and mule deer (Odocoileus hemionus) responded to the spatial context surrounding a patch. Evidence ratios of Akaike weights demonstrated that models containing effects of higher levels in the hierarchy on residence time at lower levels received up to 34 times more support in the data than models that failed to consider the higher level context for grizzly bears and up to 48 times more support for mule deer. We conclude that foraging by large herbivores is influenced by more than one level of heterogeneity in patch hierarchies and that simple empirical models offer a viable alternative to optimal foraging models for the prediction of patch residence times.     

Linking deer browsing and terpene production among genetic identities in Chamaecyparis nootkatensis and Thuja plicata (Cupressaceae)

To investigate whether differential herbivore browsing reflects genetic variation in plant defense expression, variation in needle terpenes and damage caused by black-tailed deer (Odocoileus hemionus) was analyzed on yellow-cedar (Chamaecyparis nootkatensis) and western redcedar (Thuja plicata). In a 100-genet yellow-cedar population, three genets that were heavily browsed and had extremely low levels of monoterpenes (0-0.36% dry matter), sesquiterpenes, and diterpenes were compared to unbrowsed genets (0.85-3.83% monoterpenes in dry matter). These differences were maintained in individuals protected from browsing, suggesting genetically based variation in constitutive terpene production. In western redcedar, heavily browsed trees had significantly lower total monoterpene concentrations (1.69% dry matter) than lightly browsed trees (3.32% dry matter). One heavily browsed tree expressed no monoterpenes. No differences were found for diterpenes. In both species, the genotypes with extremely low monoterpene concentrations came from the same open-pollinated families.     

Habitat selection by a large herbivore at multiple spatial and temporal scales is primarily governed by food resources

Habitat selection can be considered as a hierarchical process in which animals satisfy their habitat requirements at different ecological scales. Theory predicts that spatial and temporal scales should co‐vary in most ecological processes and that the most limiting factors should drive habitat selection at coarse ecological scales, but be less influential at finer scales. Using detailed location data on roe deer Capreolus capreolus inhabiting the Bavarian Forest National Park, Germany, we investigated habitat selection at several spatial and temporal scales. We tested 1) whether time‐varying patterns were governed by factors reported as having the largest effects on fitness, 2) whether the trade‐off between forage and predation risks differed among spatial and temporal scales and 3) if spatial and temporal scales are positively associated. We analysed the variation in habitat selection within the landscape and within home ranges at monthly intervals, with respect to land‐cover type and proxys of food and cover over seasonal and diurnal temporal scales. The fine‐scale temporal variation follows a nycthemeral cycle linked to diurnal variation in human disturbance. The large‐scale variation matches seasonal plant phenology, suggesting food resources being a greater limiting factor than lynx predation risk. The trade‐off between selection for food and cover was similar on seasonal and diurnal scale. Habitat selection at the different scales may be the consequence of the temporal variation and predictability of the limiting factors as much as its association with fitness. The landscape of fear might have less importance at the studied scale of habitat selection than generally accepted because of the predator hunting strategy. Finally, seasonal variation in habitat selection was similar at the large and small spatial scales, which may arise because of the marked philopatry of roe deer. The difference is supposed to be greater for wider ranging herbivores.     

The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions

Diet selection by mammalian herbivores is often influenced by plant community composition, and numerous studies have focused on the relationships between herbivore foraging decisions and food/plant species abundance. However, few have examined the role of neighbour palatability in affecting foraging of a target plant by large mammalian herbivores. We used a large-scale field dataset on diet selection by red deer Cervus elaphus in Fiordland National Park, New Zealand to: (1) estimate the palatability of native forest plant species to introduced deer from observed patterns of browse damage; and (2) examine whether intraspecific variation in browsing of plants can be related to variation in the local abundance of alternative forage species. Overall, 21 of the 53 forest species in our dataset were never browsed by deer. At a community level, plants were more likely to be browsed if they were in a patch of vegetation of high forage quality, containing high abundances of highly palatable species and/or low abundances of less-palatable species. Our findings suggest that deer make foraging decisions at both a coarse-grain level, selecting vegetation patches within a landscape based on the overall patch quality, and at a fine-grain level by choosing among individual plants of different species.     

The effect of terrain and female density on survival of neonatal white‐tailed deer and mule deer fawns

Juvenile survival is a highly variable life‐history trait that is critical to population growth. Antipredator tactics, including an animal's use of its physical and social environment, are critical to juvenile survival. Here, we tested the hypothesis that habitat and social characteristics influence coyote (Canis latrans) predation on white‐tailed deer (Odocoileus virginianus) and mule deer (O. hemionus) fawns in similar ways during the neonatal period. This would contrast to winter when the habitat and social characteristics that provide the most safety for each species differ. We monitored seven cohorts of white‐tailed deer and mule deer fawns at a grassland study site in Alberta, Canada. We used logistic regression and a model selection procedure to determine how habitat characteristics, climatic conditions, and female density influenced fawn survival during the first 8 weeks of life. Fawn survival improved after springs with productive vegetation (high integrated Normalized Difference Vegetation Index values). Fawns that used steeper terrain were more likely to survive. Fawns of both species had improved survival in years with higher densities of mule deer females, but not with higher densities of white‐tailed deer females, as predicted if they benefit from protection by mule deer. Our results suggest that topographical variation is a critical resource for neonates of many ungulate species, even species like white‐tailed deer that use more gentle terrain when older. Further, our results raise the possibility that neonatal white‐tailed fawns may benefit from associating with mule deer females, which may contribute to the expansion of white‐tailed deer into areas occupied by mule deer.     

Orientation behavior of Propylaea japonica toward visual and olfactory cues from its prey–host plant combination

The lady beetle Propylaea japonica (Thunberg) (Coleoptera: Coccinellidae) is an important predator of aphids in agroecosystems. The inundative release of coccinellid beetles can be an effective biological control strategy. An understanding of how biological control agents perceive and use stimuli from host plants is the key to successfully implement commercially produced predators. Here, we studied the relative role of visual and volatile cues. Dual‐choice assays using foraging‐naïve and foraging‐experienced P. japonica adults were conducted using cotton plants [Gossypium hirsutum L. (Malvaceae)] with or without infestation by the cotton aphid, Aphis gossypii (Glover) (Hemiptera: Aphididae). Overall, experienced beetles were more attracted than naïve beetles toward cues associated with aphid‐infested plants. Experienced beetles were also more responsive to olfactory cues compared with naïve beetles. Both foraging‐naïve and ‐experienced lady beetles integrate olfactory and visual cues from plants infested with aphids, with an apparently greater reliance on olfactory cues. The results suggest that foraging experience may increase prey location in P. japonica.     

Does Supplemental Feed Increase Selective Foraging in a Browsing Ungulate?

ABSTRACT Provision of supplemental feed to large herbivores is a common management practice that may motivate selective foraging, thereby influencing plant community composition. Our objective was to assess the effect of a high-quality supplement on diet composition and nutritional quality for white-tailed deer (Odocoileus virginianus). We permanently released hand-reared deer into 4 81-ha enclosures; in 2 enclosures we provided a pelleted supplement. We conducted bite-count studies seasonally to assess diet composition and quality. Supplemented deer reduced mast (fruits and pods of woody plants and cacti) in their diets (P < 0.019) during spring and autumn compared to unsupplemented deer. Diets of deer in supplemented enclosures had 2 times greater proportion of browse during spring (P = 0.065) and 5 times greater proportion of forbs during autumn (P = 0.007). Quality of the forage portion of the diet did not vary by treatment during winter or summer. Metabolizable energy concentration was 13% greater (P = 0.054) in spring and digestible protein content was 3 times greater (P = 0.006) during autumn in diets of supplemented compared to unsupplemented deer. Our results support the selective foraging hypothesis during autumn but not during winter, spring, or summer. Furthermore, white-tailed deer did not reduce the proportion of their diet composed of browse, but did reduce consumption of mast. Supplemented deer continued to eat poor-quality, chemically defended forage, perhaps to alleviate ruminal acidosis induced by the supplement or because nutrients in the supplement increased the deer's ability to detoxify chemically defended browses. A decline in mast consumption by supplemented deer could influence plant communities, depending on the role of deer in seed dispersal and seed predation. Impacts of supplemental feed on selective foraging of white-tailed deer in shrub-dominated rangelands are more complex than suggested by previous research. Long-term studies of vegetation communities are needed before wildlife managers will be able to fully incorporate effects of supplemental feed into management decisions.     

A hidden Markov model to identify and adjust for selection bias: an example involving mixed migration strategies

An important assumption in observational studies is that sampled individuals are representative of some larger study population. Yet, this assumption is often unrealistic. Notable examples include online public‐opinion polls, publication biases associated with statistically significant results, and in ecology, telemetry studies with significant habitat‐induced probabilities of missed locations. This problem can be overcome by modeling selection probabilities simultaneously with other predictor–response relationships or by weighting observations by inverse selection probabilities. We illustrate the problem and a solution when modeling mixed migration strategies of northern white‐tailed deer (Odocoileus virginianus). Captures occur on winter yards where deer migrate in response to changing environmental conditions. Yet, not all deer migrate in all years, and captures during mild years are more likely to target deer that migrate every year (i.e., obligate migrators). Characterizing deer as conditional or obligate migrators is also challenging unless deer are observed for many years and under a variety of winter conditions. We developed a hidden Markov model where the probability of capture depends on each individual's migration strategy (conditional versus obligate migrator), a partially latent variable that depends on winter severity in the year of capture. In a 15‐year study, involving 168 white‐tailed deer, the estimated probability of migrating for conditional migrators increased nonlinearly with an index of winter severity. We estimated a higher proportion of obligates in the study cohort than in the population, except during a span of 3 years surrounding back‐to‐back severe winters. These results support the hypothesis that selection biases occur as a result of capturing deer on winter yards, with the magnitude of bias depending on the severity of winter weather. Hidden Markov models offer an attractive framework for addressing selection biases due to their ability to incorporate latent variables and model direct and indirect links between state variables and capture probabilities.     

Spatial memory shapes migration and its benefits: evidence from a large herbivore

From fine‐scale foraging to broad‐scale migration, animal movement is shaped by the distribution of resources. There is mounting evidence, however, that learning and memory also guide movement. Although migratory mammals commonly track resource waves, how resource tracking and memory guide long‐distance migration has not been reconciled. We examined these hypotheses using movement data from four populations of migratory mule deer (n = 91). Spatial memory had an extraordinary influence on migration, affecting movement 2–28 times more strongly than tracking spring green‐up or autumn snow depth. Importantly, with only an ability to track resources, simulated deer were unable to recreate empirical migratory routes. In contrast, simulated deer with memory of empirical routes used those routes and obtained higher foraging benefits. For migratory terrestrial mammals, spatial memory provides knowledge of where seasonal ranges and migratory routes exist, whereas resource tracking determines when to beneficially move within those areas.     

Opposing deer and caterpillar foraging preferences may prevent reductions in songbird prey biomass in historically overbrowsed forests

Overbrowsing by ungulates decimates plant populations and reduces diversity in a variety of ecosystems, but the mechanisms by which changes to plant community composition influence other trophic levels are poorly understood. In addition to removal of avian nesting habitat, browsing is hypothesized to reduce bird density and diversity through reduction of insect prey on browse‐tolerant hosts left behind by deer. In this study, we excluded birds from branches of six tree species to quantify differences in songbird prey removal across trees that vary in deer browse preference. Early in the breeding season, birds preyed on caterpillars at levels proportional to their abundance on each host. Combining these data with tree species composition data from stands exposed to experimentally controlled deer densities over 30 years ago, we tested whether overbrowsing by white‐tailed deer reduces prey biomass long after deer densities are reduced. Our analysis predicts total prey availability in the canopy of regenerating forests is fairly robust to historic exposure to high deer densities, though distribution of prey available from host species changes dramatically. This predicted compensatory effect was unexpected and is driven by high prey abundance on a single host tree species avoided by browsing deer, Prunus serotina. Thus, while we confirm that prey abundance on host trees can act as a reliable predictor for relative prey availability, this study shows that quantifying prey abundance across host trees is essential to understanding how changes in tree species composition interact with ungulate browse preference to determine prey availability for songbirds.     

Possible use of heterospecific food-associated calls of macaques by sika deer for foraging efficiency

Heterospecific communication signals sometimes convey relevant information for animal survival. For example, animals use or eavesdrop on heterospecific alarm calls concerning common predators. Indeed, most observations have been reported regarding anti-predator strategies. Use of heterospecific signals has rarely been observed as part of a foraging strategy. Here, I report empirical evidence, collected using playback experiments, showing that Japanese sika deer, Cevus nippon, use heterospecific food calls of Japanese macaques, Macaca fuscata yakui, for foraging efficiency. The deer and macaques both inhabit the wild forest of Yakushima Island with high population densities and share many food items. Anecdotal observations suggest that deer often wait to browse fruit falls under the tree where a macaque group is foraging. Furthermore, macaques frequently produce food calls during their foraging. If deer effectively obtain fruit from the leftovers of macaques, browsing fruit fall would provide a potential benefit to the deer, and, further, deer are likely to associate macaque food calls with feeding activity. The results showed that playback of macaque food calls under trees gathered significantly more deer than silence control periods. These results suggest that deer can associate macaque food calls with foraging activities and use heterospecific calls for foraging efficiency.     

Impact of white‐tailed deer on the spread of Borrelia burgdorferi

There is a public perception that the white‐tailed deer Odocoileus virginianus (Artiodactyla: Cervidae) is the main reservoir supporting the maintenance and spread of the causative agent of Lyme disease, Borrelia burgdorferi. This study examines the pathogen prevalence rate of Borrelia in adult Ixodes scapularis (Ixodida: Ixodidae), the black‐legged tick, collected from white‐tailed deer and compares it with pathogen prevalence rates in adult ticks gathered by dragging vegetation in two contiguous counties west of the Hudson Valley in upstate New York. In both Broome and Chenango Counties, attached and unattached ticks harvested from white‐tailed deer had significantly lower prevalences of B. burgdorferi than those collected from vegetation. No attached ticks on deer (n = 148) in either county, and only 2.4 and 7.3% of unattached ticks (n = 389) in Broome and Chenango Counties, respectively, were harbouring the pathogen. This contrasts with the finding that 40.8% of ticks in Broome County and 46.8% of ticks in Chenango County collected from vegetation harboured the pathogen. These data suggest that a mechanism in white‐tailed deer may aid in clearing the pathogen from attached deer ticks, although white‐tailed deer do contribute to the spatial distribution of deer tick populations and also serve as deadend host breeding sites for ticks.     

Do Insectivorous Birds use Volatile Organic Compounds from Plants as Olfactory Foraging Cues? Three Experimental Tests

Some insectivorous birds orient towards insect‐defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect‐defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field‐based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore‐damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore‐damaged trees.     

Breeding behavior of female white‐tailed deer relative to conception: Evidence for female mate choice

Female white‐tailed deer (Odocoileus virginianus) are thought to choose between two behavioral strategies to maximize the quality of potential mates: sit and wait, characterized by concentrating activity within a restricted area, and excursive behavior, characterized by increased activity and excursions outside the home range. As movement patterns may influence conception, our goal was to examine the patterns of female white‐tailed deer movements to evaluate which breeding strategy was employed. We equipped 36 female white‐tailed deer with GPS collars from August 2013 to December 2015. We found that movement rate and probability of activity were greatest near the peak of the breeding season, and we observed increases in both metrics during the 40 days prior to estimated conception. Peak size of home range and core area occurred in the days surrounding conception. We found that 11 deer performed an excursion, ranging from 43 days before until 36 days after conception, with the peak probability of being outside of an individual home range occurring 1 day prior to conception. Our results suggest that female white‐tailed deer may attempt to maximize the quality of their mates by advertising availability for breeding through excursive behaviors just prior to entering estrus.     

Influence of aphid honeydew on the foraging behaviour of Hippodamia convergens larvae

1. Environmental cues associated with prey are known to increase predator foraging efficiency. Ladybird larvae are major predators of aphids. The sugary excretion of aphids (honeydew) has been proposed to serve as a prey‐associated cue for ladybird larvae. 2. Ladybird larvae are regularly found on the ground moving between plants or after falling off plants. The use of prey‐associated cues would be particularly beneficial for ladybird larvae on the ground in that such cues would help them to decide which plants to climb because aphids are patchily distributed within as well as amongst plants and, as a result, many plants are either not infested with aphids or do not host an aphid species of high nutritional value for ladybird larvae. 3. Laboratory experiments with larvae of Hippodamia convergens Guérin‐Méneville (Coleoptera: Coccinellidae) were carried out to explore whether honeydew accumulated on the ground is used as a foraging cue. The study also investigated whether, if honeydew is a foraging cue, larvae show differential responses to honeydew of high‐quality prey Acyrthosiphon pisum Harris compared with that of low‐quality prey Aphis fabae Scopoli (both: Homoptera: Aphididae). 4. Hippodamia convergens larvae stayed longer in areas containing honeydew but did not engage in longer bouts of searching. Furthermore, larvae did not distinguish between honeydew from high‐ and low‐quality aphid prey.     

Visual cues override olfactory cues in the host-finding process of the monophagous leaf beetle Altica engstroemi

It is generally assumed that specialist insect herbivores utilize plant odours to find their particular host plants and that visual cues are of minor importance in the host‐finding process. We performed Y‐tube olfactometer bioassays and small‐scale field experiments to determine whether, under laboratory and field conditions, the monophagous herbivore Altica engstroemi J. Sahlberg (Coleoptera: Chrysomelidae: Alticinae) is guided to its host plant Filipendula ulmaria (L.) Maxim. (Rosaceae) by visual or olfactory cues. The olfactometer tests showed that A. engstroemi was never attracted to odours, either from undamaged or from damaged plants. Even starvation for 24 h did not change this behaviour. However, the field experiment showed that visual cues alone were sufficient to attract a significant number of starved beetles when offered a choice between bagged host plants and bagged green plastic control ‘plants’. Our findings contrast with the general view that plant odours constitute the major cue in the host‐finding process among specialized phytophagous insects. A review of the literature for the period 1986–2006 inclusive, relating to host‐plant finding in Chrysomelidae, identified studies of 19 chrysomelid species, all of which were guided by olfactory cues. No species were guided to their host by visual cues. Although some studies demonstrated that chrysomelids may exhibit orientation responses to colour or contrast, our study on A. engstroemi is the only one demonstrating that visual cues affect host‐plant selection in a chrysomelid species. We suggest that the use of visual cues in host‐finding may evolve among chrysomelids with limited dispersal ability in persistent habitats and may be found among species monophagous on abundant host plants that dominate the structure of the plant community, that is, where the host plant's presence is predictable in time and space.     

When perception reflects reality: Non‐native grass invasion alters small mammal risk landscapes and survival

Modification of habitat structure due to invasive plants can alter the risk landscape for wildlife by, for example, changing the quality or availability of refuge habitat. Whether perceived risk corresponds with actual fitness outcomes, however, remains an important open question. We simultaneously measured how habitat changes due to a common invasive grass (cheatgrass, Bromus tectorum) affected the perceived risk, habitat selection, and apparent survival of a small mammal, enabling us to assess how well perceived risk influenced important behaviors and reflected actual risk. We measured perceived risk by nocturnal rodents using a giving‐up density foraging experiment with paired shrub (safe) and open (risky) foraging trays in cheatgrass and native habitats. We also evaluated microhabitat selection across a cheatgrass gradient as an additional assay of perceived risk and behavioral responses for deer mice (Peromyscus maniculatus) at two spatial scales of habitat availability. Finally, we used mark‐recapture analysis to quantify deer mouse apparent survival across a cheatgrass gradient while accounting for detection probability and other habitat features. In the foraging experiment, shrubs were more important as protective cover in cheatgrass‐dominated habitats, suggesting that cheatgrass increased perceived predation risk. Additionally, deer mice avoided cheatgrass and selected shrubs, and marginally avoided native grass, at two spatial scales. Deer mouse apparent survival varied with a cheatgrass–shrub interaction, corresponding with our foraging experiment results, and providing a rare example of a native plant mediating the effects of an invasive plant on wildlife. By synthesizing the results of three individual lines of evidence (foraging behavior, habitat selection, and apparent survival), we provide a rare example of linkage between behavioral responses of animals indicative of perceived predation risk and actual fitness outcomes. Moreover, our results suggest that exotic grass invasions can influence wildlife populations by altering risk landscapes and survival.     

The effects of habitat manipulation on population distribution and foraging behavior in meadow voles

Individuals, free to choose between different habitat patches, should settle among them such that fitness is equalized. Alternatives to this ideal free distribution result into fitness differences among the patches. The concordance between fitnesses and foraging costs among inhabitants of different quality patches, demonstrated in recent studies, suggests that the mode of habitat selection and the resulting fitness patterns may have important implications to the resource use of a forager and to the survival of its prey. We studied how coarse scale selection between habitat patches of different quality and quitting harvest rate in these patches are related to each other and to fine scale patch use in meadow voles (Microtus pennsylvanicus). To demonstrate these relationships, we manipulated habitat patches within large field enclosures by mowing vegetative cover and adding supplemental food according to a 2×2 factorial design. We tracked vole population densities, collected giving‐up densities (GUDs, a measure of patch quitting harvest rate), and monitored the removal of seeds from lattice grids with 1.5 m intervals (an index of fine‐scale space use) in the manipulated habitat patches. Changes in habitat quality induced changes in habitat use at different spatial scales. In preferred habitats with intact cover, voles were despotic and GUDs were low, but increased with the addition of food. In contrast, voles in less‐preferred mowed habitats settled into an ideal free distribution, GUDs were high and uninfluenced by the addition of food. Seed removal was enhanced by the presence of cover but inhibited by supplemental food. Across all treatments, vole densities and GUDs were strongly correlated making it impossible to separate their effects on seed removal rates. However, this relationship broke down in unmowed habitats, where GUDs rather than vole density primarily influenced seed removal by voles. GUDs and seed removal correlated with predation on tree seedlings formerly planted into the enclosures, demonstrating the mechanisms between coarse‐scale habitat manipulations and community level consequences on a forager's prey.