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Sagebrush ecosystems (Artemisia spp.) face many threats including large wildfires and conversion to invasive annuals, and thus are the focus of intense restoration efforts across the western United States. Specific attention has been given to restoration of sagebrush systems for threatened herbivores, such as Greater Sage-Grouse (Centrocercus urophasianus) and pygmy rabbits (Brachylagus idahoensis), reliant on sagebrush as forage. Despite this, plant chemistry (e.g., crude protein, monoterpenes and phenolics) is rarely considered during reseeding efforts or when deciding which areas to conserve. Near-infrared spectroscopy (NIRS) has proven effective in predicting plant chemistry under laboratory conditions in a variety of ecosystems, including the sagebrush steppe. Our objectives were to demonstrate the scalability of these models from the laboratory to the field, and in the air with a hyperspectral sensor on an unoccupied aerial system (UAS). Sagebrush leaf samples were collected at a study site in eastern Idaho, USA. Plants were scanned with an ASD FieldSpec 4 spectroradiometer in the field and laboratory, and a subset of the same plants were imaged with a SteadiDrone Hexacopter UAS equipped with a Rikola hyperspectral sensor (HSI). All three sensors generated spectral patterns that were distinct among species and morphotypes of sagebrush at specific wavelengths. Lab-based NIRS was accurate for predicting crude protein and total monoterpenes (R2 = 0.7–0.8), but the same NIRS sensor in the field was unable to predict either crude protein or total monoterpenes (R2 2&thinsp

The amount and composition of phytochemicals in forage plants influences habitat quality for wild herbivores. However, evaluating forage quality at fine resolutions across broad spatial extents (i.e., foodscapes) is challenging. Unmanned aerial systems (UAS) provide an avenue for bridging this gap in spatial scale. We evaluated the potential for UAS technology to accurately predict nutritional quality of sagebrush (Artemisia spp.) across landscapes. We mapped seasonal forage quality across two sites in Idaho, USA, with different mixtures of species but similar structural morphotypes of sagebrush. We classified the sagebrush at both study sites using structural features of shrubs with object-based image analysis and machine learning and linked this classification to field measurements of phytochemicals to interpolate a foodscape for each phytochemical with regression kriging. We compared fine-scale landscape patterns of phytochemicals between sites and seasons. Classification accuracy for morphotypes was high at both study sites (81–87%). Forage quality was highly variable both within and among sagebrush morphotypes. Coumarins were the most accurately mapped (r2 = 0.57–0.81), whereas monoterpenes were the most variable and least explained. Patches with higher crude protein were larger and more connected in summer than in winter. UAS allowed for a rapid collection of imagery for mapping foodscapes based on the phytochemical composition of sagebrush at fine scales but relatively broad extents. However, results suggest that a more advanced sensor (e.g., hyperspectral camera) is needed to map mixed species of sagebrush or to directly measure forage quality.

Large wild herbivores are important and natural components of forest ecosystems, but through their browsing activities have the potential to influence the structure and composition of forest communities, thus timber production and ecosystem dynamics. To examine the effects of browsing by wild herbivores on a young post-disturbance forest in the Kysuce region of northwestern Slovakia, we established two sets of 2 m radius plots, 15 within a fenced area (5.12 ha) that excluded large wild herbivores, and 15 within an adjacent unfenced area. In each plot, we recorded the species, tree height, stem base diameter, and mutual geographic positions of trees. When we compared tree community characteristics between the unfenced and fenced plots, we found fewer and smaller broadleaved tree species, except silver birch (Betula pendula Roth.) in the unfenced plots. Although common rowan (Sorbus aucuparia L.) was the dominant species within fenced plots, where some individuals were over 6.0 m tall, this species was rare outside the fenced area and usually did not exceed 1.5 m. In contrast, Norway spruce (Picea abies Karts L.) was more abundant and taller within the unfenced area, likely released from competition by suppression of broadleaved trees by herbivores. In addition, fenced plots also showed twice the tree species richness (Shannon index) of unfenced ones. Despite changes in tree communities, total aboveground biomass stock was only slightly but significantly lower in the unfenced than the fenced plots (29.6 kg per 10 m2 vs. 33.5 kg per 10 m2). Our study suggested that browsing pressure by large wild herbivores that focused on most broadleaved trees weakened interspecies competition and allowed the expansion of Norway spruce. As a consequence, converting spruce monocultures to mixed species stands is likely unrealistic when faced with heavy browsing pressure by wild large herbivores.

In heterogeneous landscapes, large herbivores employ plastic behavioral strategies to buffer themselves against negative effects of environmental variation on fitness. Yet, the mechanisms by which individual responses to such variation scale up to influence population performance remain uncertain. Analyses of space-use behaviors exemplify this knowledge gap, because such behaviors are often assumed, but rarely demonstrated, to have direct fitness consequences. We combined fine-scale data on forage biomass and quality with movement data and measures of somatic energy reserves to determine whether variation in use (the quantity of resource units, e.g., pixels on a landscape, that receive some level of investment by an animal during a specific sampling period) or selection (use of a resource unit relative to its availability to the animal during the same sampling period) of the nutritional landscape predicted early winter body condition of mule deer (Odocoileus hemionus). At the population level, mule deer exhibited stronger selection for high forage biomass at the landscape scale than at the home-range scale, and during summer than during spring. Use of the nutritional landscape varied among individual deer and had important consequences for early winter condition (an important determinant of survival and reproduction in capital-breeding ungulates). Females that consistently used vegetation communities that provided high biomass of preferred forage plants throughout spring and summer entered winter in better condition than females that used those vegetation communities less frequently. In contrast, selection (i.e., use relative to availability) of the nutritional landscape by individual deer was not significantly related to early winter condition at either the landscape or home-range scales. Our results highlight the value of using mechanistic, nutritional approaches to understand the potential fitness consequences of individual variation in behavior. In addition, our study suggests that patterns of forage use by ungulates may sometimes correlate more strongly with fitness than patterns of forage selection, which are scale-dependent and more vulnerable to biases stemming from the need to accurately quantify availability.

Herbivores that forage on chemically defended plants consume complex mixtures of plant secondary metabolites (PSMs). However, the mechanisms by which herbivores tolerate mixtures of PSMs are relatively poorly understood. As such, it remains difficult to predict how PSMs, singly or as complex mixtures, influence diet selection by herbivores. Although relative rates of detoxification of PSMs have been used to explain tolerance of PSMs by dietary specialist herbivores, few studies have used the rate of detoxification of individual PSMs to understand dietary preferences of individual herbivores for individual versus mixtures of PSMs. We coupled in vivo experiments using captive feeding trials with in vitro experiments using enzymatic detoxification assays to evaluate the dietary preferences and detoxification capacities of pygmy rabbits (Brachylagus idahoensis), dietary specialists on sagebrush (Artemisia spp.), and mountain cottontails (Sylvilagus nuttallii), dietary generalists. We compared preference for five single PSMs in sagebrush compared to a mixture containing those same five PSMs. We hypothesized that relative preference for individual PSMs would coincide with faster detoxification capacity for those PSMs by specialists and generalists. Pygmy rabbits generally showed little preference among individual PSMs compared to mixed PSMs, whereas mountain cottontails exhibited stronger preferences. Pygmy rabbits had faster detoxification capacities for all PSMs and consumed higher concentrations of individual PSMs versus a mixture than cottontails. However, detoxification capacity for an individual PSM did not generally coincide with preferences or avoidance of individual PSMs by either species. Cottontails avoided, but pygmy rabbits preferred, camphor, the PSM with the slowest detoxification rate by both species. Both species avoided β-pinene despite it having one of the fastest detoxification rate. Taken together our in vivo and in vitro results add to existing evidence that detoxification capacity is higher in dietary specialist than generalist herbivores. However, results also suggest that alternative mechanisms such as absorption and the pharmacological action of individual or mixtures of PSMs may play a role in determining preference of PSMs within herbivore species.

Large herbivores prefer to consume plants with high concentration of digestible carbohydrates and protein, whereas fire spread is favored by foliage characteristics that induce rapid ignition and high radiant energy. We tested in eight Patagonian woody plants whether foliar traits that determine dry matter and nitrogen digestibility to browsing herbivores were related to four variables that determine their flammability. We found a negative relation between digestibility and flammability. Fiber content (i.e. Neutral and Acid Detergent Fiber), was positively related with heat emission, maximum temperature and flame duration, being negatively related to time to ignition. PCA indicated that digestibility was negatively related with all variables associated with flammability. Digestibility decreased throughout the growing season, whereas flammability showed no clear seasonal trend. Fiber content was correlated with time to ignition and maximum temperature during most of the growing season, except in late spring. We conclude that highly digestible plants are likely to be less flammable than fiber rich ones, and that fiber content may serve as an indicator of seasonal changes in flammability of foliage. Because leaf traits are likely to influence habitat selection and movement by herbivores as well as spread and severity of fires, further studies linking foliar shape variables and plant architecture with their chemical composition may improve current mechanistic models of herbivore and fire behavior.

Forested lands in the western USA have undergone changes in management and condition that are resulting in a shift towards climax vegetation. These changes can influence the quality and quantity of forage for herbivores that rely on early-seral plants. To evaluate how management of forested landscapes might affect nutrition for Shiras moose (A. a. shirasi) at large spatial scales, we focused on shrubs and evaluated summer diet composition, forage availability, and forage quality across 21 population management units encompassing >36,000 km2 in northern Idaho, USA. We identified 17 shrub species in the diets of moose, 11 of which comprised the bulk of the diets. These forage shrubs varied markedly in both energy (mean digestible energy for leaves ranged from 9.62 to 12.89 kJ/g) and protein (mean digestible protein for leaves ranged from 1.73 to 7.90%). By adapting established field sampling methods and integrating recent advances in remote sensing analyses in a modeling framework, we predicted approximations of current and past (i.e., 1984) quantities of forage shrubs across northern Idaho. We also created a qualitative index of population trend for moose across population management units using harvest data. Predicted quantities of forage shrubs varied widely across the study area with generally higher values at more northern latitudes. The quantity of forage shrubs was estimated to have declined over the past 30 years in about half of the population management units, with the greatest declines predicted for high-energy forage species. The population trend index was correlated with the percent change in availability of moderate-energy forage shrubs, indicating that availability of forage shrubs and change in availability over time might be affecting population dynamics for moose in northern Idaho. Our study highlights the importance of assessing how changes in forest management across broad spatiotemporal extents could affect wildlife and their habitats.

Sagebrush ( Artemisia spp.) habitat in the Intermountain West is one of the most endangered ecosystems in North America due, in part, to fire, climate change, and anthropogenic disturbances. However, restoration efforts rarely consider the dietary quality of sagebrush that is conserved or restored despite growing evidence that it is an influential parameter explaining habitat use by many important wild and domestic herbivores. The objective of this study was to evaluate the capacity of near-infrared reflectance spectroscopy (NIRS) to measure and monitor the dietary quality of sagebrush. Leaf samples were collected from two sagebrush species over two seasons and three sites in Idaho, USA. We developed calibration equations for crude protein (CP), dry matter digestibility (DMD), 1,8-cineole (cineole), and total polyphenolics. The coefficient of determination ( r 2 ) and ratio of performance to deviation (RPD) were 0.93 and 3.5 for CP, 0.83 and 1.8 for DMD, 0.64 and 1.5 for cineole, and 0.64 and 1.6 for total polyphenolics. These results indicate that NIRS may offer a rapid, noninvasive, diagnostic tool for assessing dietary quality of sagebrush, but future research should explore the potential for development of improved prediction equations and in situ analysis of sagebrush dietary quality with field spectroscopy.

Understanding habitat use by animals requires understanding the simultaneous tradeoffs between food and predation risk within a landscape. Quantifying the synergy between patches that provide quality food and those that are safe from predators at a scale relevant to a foraging animal could better reveal the parameters that influence habitat selection. To understand more thoroughly how animals select habitat components, we investigated tradeoffs between diet quality and predation risk in a species endemic to sagebrush Artemisia spp. communities in North America, the pygmy rabbit Brachylagus idahoensis. This species is a rare example of a specialist herbivore that relies almost entirely on sagebrush for food and cover. We hypothesized that pygmy rabbits would forage in areas with low food risk (free of plant secondary metabolites, PSMs) and low predation risk (high concealment). However, because of relatively high tolerance to PSMs in sagebrush by pygmy rabbits, we hypothesized that they would trade off the...

When selecting habitats, herbivores must weigh multiple risks, such as predation, starvation, toxicity, and thermal stress, forcing them to make fitness trade-offs. Here, we applied the method of paired comparisons (PC) to investigate how herbivores make trade-offs between habitat features that influence selection of food patches. The method of PC measures utility and the inverse of utility, relative risk, and makes trade-offs and indifferences explicit by forcing animals to make choices between two patches with different types of risks. Using a series of paired-choice experiments to titrate the equivalence curve and find the marginal rate of substitution for one risk over the other, we evaluated how toxin-tolerant (pygmy rabbit Brachylagus idahoensis) and fiber-tolerant (mountain cottontail rabbit Sylviagus nuttallii) herbivores differed in their hypothesized perceived risk of fiber and toxins in food. Pygmy rabbits were willing to consume nearly five times more of the toxin 1,8-cineole in their diets to avoid consuming higher levels of fiber than were mountain cottontails. Fiber posed a greater relative risk for pygmy rabbits than cottontails and cineole a greater risk for cottontails than pygmy rabbits. Our flexible modeling approach can be used to (1) quantify how animals evaluate and trade off multiple habitat attributes when the benefits and risks are difficult to quantify, and (2) integrate diverse risks that influence fitness and habitat selection into a single index of habitat value. This index potentially could be applied to landscapes to predict habitat selection across several scales.

When animals select habitats, they integrate a suite of behaviors that are influenced by multiple competing resource requirements. Resources that influence decisions about habitat use are likely to differ across spatial scales and hierarchical behaviors. At coarser scales, animals are expected to select resources that are critical to fitness, and at finer scales, to intensively use resources that enhance fitness. Our goal was to contrast habitat selection at two hierarchical behavioral levels (patch selection and intensity of patch use) to test hypotheses about how resources shape habitat use. We applied a two-stage hurdle model to quantify both initial selection and intensity of use of resource patches by a burrowing herbivore, the pygmy rabbit (Brachylagus idahoensis). We expected security from predation to influence initial selection of patches, and forage availability to influence intensity of use of selected patches. We monitored locations of adults fitted with radio-collars during winter and summer. We measured vegetation, burrow characteristics, and concealment cover within patches that were used and unused by rabbits, and we quantified burrow densities surrounding patches in a GIS. Selection of used patches from available patches was largely influenced by security resources (presence and proximity of burrows, shrub height, and woody ground cover) during both seasons. Intensity of patch use was also influenced by forage availability and, consequently, differed between seasons. During winter, sagebrush is the primary forage, and greater sagebrush canopy within a patch was associated with increased intensity of use. During summer, rabbits more intensively used patches with greater availability of herbaceous forage. Elucidating how animals make choices about habitats across a diversity of spatial and temporal scales will continue to increase our understanding of the factors that govern distribution of populations and movements of individuals. By extending these concepts to hierarchical behaviors, we can enhance insights into the processes that shape the patterns of habitat use we observe.

Both temperature and diet quality play an important role in the time and energy budgets of small mammalian herbivores. However, little is known about how temperature and diet quality interact to influence diet selection, nutrient extraction, and energetics, and how these effects might differ among species. Therefore, we examined the effects of diet quality and temperature on aspects of the foraging ecology of two species of lagomorphs, pygmy rabbits (Brachylagus idahoensis), which are small dietary specialists, and mountain cottontail rabbits (Sylvilagus nuttallii), which are larger dietary and habitat generalists. In a series of feeding experiments, we investigated 1) the effects of temperature on selection of plant fiber and the plant secondary metabolite 1,8 cineole in their diets, 2) effects of temperature and plant fiber on daily intake, digestion, and passage of food, 3) effects of plant fiber and 1,8 cineole on resting metabolic rate, and 4) how these interactions differ between the rabbit species. Both species chose to eat more total food and a greater proportion of high fiber food that passed more quickly through the digestive system in colder temperatures. However, temperature did not affect how much 1,8 cineole the rabbits consumed nor how thoroughly they digested food. Food quality affected how well they digested the dry matter in the food, but not their resting metabolic rate. Understanding how the interactions between ambient temperature and food quality affect selection of diets and intake by small mammalian herbivores, and the physiological mechanisms governing these choices, is useful for predicting how these species might respond to changes in both temperature and food quality and inform conservation and restoration strategies.

For herbivores, nutrient intake is limited by the relatively low nutritional quality of plants and high concentrations of potentially toxic defensive compounds (plant secondary metabolites [PSMs]) produced by many plants. In response to phytochemical challenges, some herbivores selectively forage on plants with higher nutrient and lower PSM concentrations relative to other plants. Pygmy rabbits (Brachylagus idahoensis) are dietary specialists that feed on sagebrush (Artemisia spp.) and forage on specific plants more than others within a foraging patch. We predicted that the plants with evidence of heavy foraging (browsed plants) would be of higher dietary quality than plants that were not browsed (unbrowsed). We used model selection to determine which phytochemical variables best explained the difference between browsed and unbrowsed plants. Higher crude protein increased the odds that plants would be browsed by pygmy rabbits and the opposite was the case for certain PSMs. Additionally, because pygmy rabbits can occupy foraging patches (burrows) for consecutive years, their browsing may influence the nutritional and PSM constituents of plants at the burrows. In a post hoc analysis, we did not find a significant relationship between phytochemical concentrations, browse status, and burrow occupancy length. We concluded that pygmy rabbits use nutritional and chemical cues while making foraging decisions.

Animals face multiple risks while foraging such as the risk of acquiring inadequate energy from food and the risk of predation. We evaluated how two sympatric rabbits (pygmy rabbits, Brachylagus idahoensis, and mountain cottontail rabbits, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush-steppe of western North America respond to different types and levels of perceived risks (i.e., fitness cost × probability of occurrence), including fiber and toxins in food, exposure to predation, and distance from a refuge. We measured food intake by the rabbits at paired food patches that varied in these risks and used the method of paired comparisons to create a relative ranking of habitat cues, which revealed an animal’s perceived risk on a single scale representing an integrated response to a variety of risks. Pygmy rabbits perceived exposure to predation risk and distance from a burrow as riskier than did cottontails, whereas cottontails perceived dietary toxin as riskier. Pygmy rabbits consumed lower quality food, containing higher fiber or toxins, thereby avoided feeding in exposed patches or traveling far from their burrow to forage. In contrast, cottontails fed in exposed patches and traveled farther from the burrow to obtain higher quality food. We have shown how risks can be integrated into a single model that allows animals to reveal their perceptions of risks on a single scale that can be used to create a spatially explicit landscape of risk.

Organisms respond to their heterogeneous environment in complex ways at many temporal and spatial scales. Here, I examine how the smallest scale process in foraging by mammalian herbivores, taking a bite, influences plants and herbivores over larger scales. First, because cropping bites competes with chewing them, bite size influences short-term intake rate of herbivores within plant patches. On the other hand, herbivores can chew bites while searching for new ones, thus influencing the time spent vigilant and intake rate as animals move among food patches. Therefore, bite size affects how much time herbivores must spend foraging each day. Because acquiring energy is necessary for fitness, herbivores recognize the importance of bite size and select bites, patches and diets based on tradeoffs between harvesting rates, digestion, and sheering forces. In turn, induced structural defenses of plants, such as thorns, allow plants to respond immediately to herbivory by reducing bite size and thus tissue loss. Over evolutionary time, herbivores have adapted mouth morphology that allows them to maximize bite size on their primary forage plant, whereas plants faced with large mammalian herbivores have adapted structures such as divarication that minimize bite size and protect themselves from herbivory. Finally, bite size available among plant communities can drive habitat segregation and migration of larger herbivores across landscapes.

The question of how much time a foraging herbivore should spend in a patch of food poses a central challenge in classical foraging theory. However, there remains uncertainty about the relevance of the patch paradigm to foraging decisions by large herbivores. This paper examines evidence for successfully predicting and quantifying patch departure decisions for large mammalian herbivores foraging across several spatial and temporal scales. Departure decisions at fine scales are influenced by tradeoffs between maximizing intake rate and food quality. Classical models for departure decisions at larger spatial scales, particularly the marginal value theorem, appear inadequate. We advocate exploring alternative models for predictions of residence time at the patch scale.

Summary 1 The gain function describes the amount of food consumed in a patch as a function of patch residence time. Gain functions play a central role in foraging theory but alternative functional forms portraying dynamics of gain have not been evaluated. We evaluated the strength of evidence in the data for alternative gain functions of mule deer (Odocoileus hemionus, Rafinesque 1817) and blue duikers (Cephalophus monticola, Blythe 1848) feeding in patches composed of different plant species and plant sizes. 2 Gain functions decelerated with patch residence time, but there was considerable variation among individual animals and patch types in the nature of this response. Asymptotic and piecewise-linear models received the greatest support in the data. 3 Deceleration in gain was caused by a composite of effects that retarded instantaneous intake rate, including reductions in bite mass and increases in bite interval (time between successive bites). Bite interval increased as a result of increases in processing time of accumulated forage in the mouth, rather than increases in time allocated to cropping. 4 We demonstrated that unwarranted assumptions about the shape of gain functions can have profound effects on predictions of patch models. Predictions of the classical patch model using purely asymptotic gain functions contrasted sharply with predictions of model-averaged gain functions that were supported by the data.

The carbon-isotope composition of hair and feces offers a glimpse into the diets of mammalian herbivores. It is particularly useful for determining the relative consumption of browse and graze in tropical environments, as these foods have strongly divergent carbon-isotope compositions. Fecal δ13C values reflect the last few days consumption, whereas hair provides longer term dietary information. Previous studies have shown, however, that some fractionation occurs between dietary δ13C values and those of hair and feces. Accurate dietary reconstruction requires an understanding of these fractionations, but few controlled-feeding studies have been undertaken to investigate these fractionations in any mammalian taxa, fewer still in large mammalian herbivores. Here, we present data from the first study of carbon-isotope fractionation between diet, hair, and feces in multiple herbivore taxa. All taxa were fed pure alfalfa (Medicago sativa) diets for a minimum period of 6 months, at which point recently grown hair was shaved and analyzed for carbon isotopes. The mean observed diet–hair fractionation was +3.2‰, with a range of +2.7 to +3.5‰. We also examined diet–feces fractionation for herbivores on alfalfa and bermudagrass (Cynodon dactylon) feeds. The mean diet–feces fractionation for both diets was –0.8‰, with less fractionation for alfalfa (–0.6‰) than bermudagrass (–1.0‰). Fecal carbon turnover also varies greatly between taxa. When diets were switched, horse (Equus caballus) feces reflected the new diet within 60 h, but alpaca (Lama pacos) feces did not equilibrate with the new diet for nearly 200 h. Thus, fecal carbon isotopes provide far greater dietary resolution for hindgut-fermenting horses than foregut-fermenting alpacas.

Current land-use changes in eastern deciduous forests, such as fragmentation, may affect population sizes of native wildlife that may exacerbate declines in rare and endangered wildflower populations in the eastern deciduous forests. In this study, we examined the influence of herbivory by rodents (Peromyscus leucopus, Sciurus sp., and Tamias striatus) and white-tailed deer (Odocoileus virginianus) on the growth and survival of Turk's cap lily (Lilium superbum) planted in Virginia. Bulbs were planted in pairs and in patches. One plant per pair was protected from deer damage with a wire cage, and patches consisted of two, 10 and 25 bulbs planted within 0.04 ha. Rodents dug up and consumed 9% of all the bulbs planted, and fatal rodent damage was 3 times greater in successional than in upland hardwood and creek bottom habitats. White-tailed deer consumed the apical meristem of 28% of the unprotected lilies that emerged, reducing mean plant height and stopping growth and reproduction for that season. Deer and insects, but not rodents, damaged a greater proportion of plants emerging in small patches (1–2 plants/0.04 ha) than on larger patches (3–20 plants/0.04 ha). Therefore, when protecting remaining populations or restoring new populations of rare perennial wildflowers in the eastern deciduous forest, methods for protecting plants from herbivory by rodents and white-tailed deer should be considered.

Although well known as carnivores and not capable of digesting plant fiber, grizzly bears (Ursus arctos horribilis) consume over 200 species of plants and are entirely vegetarian in some ecosystems. Even in ecosystems with abundant meat resources, green vegetation can be an important seasonal food resource. Therefore, we examined the morphological, physiological, and environmental constraints that determine the nutritional value of herbaceous vegetation to grizzly bears. Short-term, board foraging trials were used with captive grizzly bears to determine constraints on intake rate including bite size, bite rate, bear size, plant species, plant height, and plant distribution. Feeding trials were conducted to determine the effect of protein level (12-35%) and digestible dry matter intake on weight gain. Finally, maximum daily intake, daily foraging time, and weight change were measured for captive bears foraging on highly abundant and nutritious forbs and grasses during 12-day trials. Intake during short-term board trials overestimated the intake of freely foraging bears from two- to seven-fold depending on bear size. Because of their relatively larger bite sizes, smaller absolute energy requirements, and relatively larger intake capacity, smaller bears (120 kg) made greater weight gains than very large bears on herbaceous vegetation. Smaller bears with ad libitum access to palatable, nutritious forbs gained weight at rates equal to wild bears. However, depending upon plant characteristics, bite sizes and available daily foraging time increasingly prevented large bears (120 kg) from gaining weight on herbaceous vegetation. Both captive and wild bears select forbs over grasses at similar growth stages because forbs are generally higher in protein and more digestible than grasses. Therefore, the nutritional well-being of wild grizzly bears could be improved in areas where bears are largely herbivorous and, thus, relatively small by purposefully managing for nutritious forbs.

Understanding how large herbivores select their diet requires accurate measurements of the nutritional costs and benefits of food items and the composition of plants in the diet relative to the composition of plants in the habitat. We followed moose (Alces alces) tracks in the snow and measured bites of browse plants taken and the number and mass of twigs of the 10 browse species available within the moose's reach and measured morphological and chemical characteristics of browse plants. The diets of 41 moose in northern coastal Sweden contained 75% Scots pine (Pinus sylvestris) and willow (Salix spp.), with the remaining proportion equally divided among five deciduous browse species and common juniper (Juniperus communis). The moose used approximately 20% of the bites available along the foraging patch, usually taking only one or two bites on each plant selected for browsing. Moose selected rowan (Sorbus aucaparia), willow species, and aspen (Populus tremula) in winter, used gray alder (Alnus incana), silver birch (Betula pendula), and Scots pine in proportion to their availability, and avoided pubescent birch (Betula pubescens) and Norway spruce (Picea abies). Selection was correlated with plant morphology but not with digestibility or nitrogen, fibre, or phenolic content. Moose selected browse species offering fewer, larger stems over those with many, smaller stems. Moose made slight adjustments in their food selection and foraging intensity as abundance of browse plants declined, thereby presumably maintaining higher harvesting rates.

We examined the functional response and foraging behavior of young moose (Alces alces) and white-tailed deer (Odocoileus virginianus) relative to animal size and the size and distribution of browse patches. The animals were offered one, three, or nine stems of dormant red maple (Acer rubrum) in hand-assembled patches spaced 2.33, 7, 14, or 21 m apart along a runway. Moose took larger twig diameters and bites and had greater dry matter and digestible energy intake rates than did deer, but had lower cropping rates. Moose and deer travelled at similar velocities between patches and took similar numbers of bites per stem. We found that a model of intake rate, based on the mechanics of cropping, chewing, and encountering bites, effectively described the intake rate of moose and deer feeding in heterogeneous distributions of browses. As patch size and density declined, the animals walked faster between patches, cropped larger bites, and cropped more bites per stem, and hence, dry matter intake rates remained relatively constant. As is characteristic of many hardwood browse stems, however, potential digestible energy concentration of the red maple stems declined as the size and number of bites removed (i.e., stem diameter at point of clipping) by the animals increased. Therefore, the digestible energy content of the diet declined with decreasing patch size and density. Time spent foraging within a patch increased as patch size increased and as distance between patches increased, which qualitatively supported the marginal-value theorem. However, actual patch residence times for deer and moose exceeded those predicted by the marginal-value theorem (MVT) by approximately 250%. The difference between actual and predicted residence time may have been a result of (1) an unknown or complex gain function, (2) the artificial conditions of the experiments, or (3) assumptions of MVT that do not apply to herbivores.

Pygmy rabbits (Brachylagus idahoensis) are one of only three vertebrates that subsist virtually exclusively on sagebrush (Artemisia spp.), which contains high levels of monoterpenes that can be toxic. We examined the mechanisms used by specialist pygmy rabbits to eliminate 1,8-cineole, a monoterpene of sagebrush, and compared them with those of cottontail rabbits (Sylvilagus nuttalli), a generalist herbivore. Rabbits were offered food pellets with increasing concentrations of cineole, and we measured voluntary intake and excretion of cineole metabolites in feces and urine. We expected pygmy rabbits to consume more, but excrete cineole more rapidly by using less-energetically expensive methods of detoxification than cottontails. Pygmy rabbits consumed 3–5 times more cineole than cottontails relative to their metabolic body mass, and excreted up to 2 times more cineole metabolites in their urine than did cottontails. Urinary metabolites excreted by pygmy rabbits were 20 % more highly-oxidized and 6 times less-conjugated than those of cottontails. Twenty percent of all cineole metabolites recovered from pygmy rabbits were in feces, whereas cottontails did not excrete fecal metabolites. When compared to other mammals that consume cineole, pygmy rabbits voluntarily consumed more, and excreted more cineole metabolites in feces, but they excreted less oxidized and more conjugated cineole metabolites in urine. Pygmy rabbits seem to have a greater capacity to minimize systemic exposure to cineole than do cottontails, and other cineole-consumers, by minimizing absorption and maximizing detoxification of ingested cineole. However, mechanisms that lower systemic exposure to cineole may come with a higher energetic cost in pygmy rabbits than in other mammalian herbivores.

The rate of food intake exerts an important influence on many aspects of herbivore ecology, including diet and habitat choices, social organization, and predator avoidance. When food is spatially concentrated, short-term dry matter intake rate (I, g/min) is determined largely by morphology of the mouth and mechanics of food consumption. Morphology (tooth size and jaw musculature) and mechanics (cropping and chewing processes) are hypothesized to scale with body mass (M) for mammalian herbivores. By using a simple model of processes regulating short-term I, we developed and tested hypotheses on the scaling of these parameters in 12 species of mammalian herbivores whose masses ranged from 0.05 kg to 547 kg. Specifically, this model predicts that I is controlled by the size of bite taken, by the time required to crop a bite, and by the rate at which food in the mouth can be processed. Maximum bite size scaled with M0 72, whereas cropping time did not scale with body mass and averaged 0.015 min/bite across sp...

Understanding dietary specialization in herbivores has theoretical and practical implications in ecology, yet defining niche breadth consistently has been problematic. To increase clarity and communication among ecologists and among disciplines (i.e., chemists, pharmacologists), we propose a specialization key for mammalian herbivores that assigns "obligatory" and "facultative" modifiers to the terms "specialist" and "generalist". These modifiers are assigned based on (1) relative breadth of the animal's realized niche and diet (what it eats), (2) relative breadth of the fundamental niche and available diet (what it could eat), (3) the extent of chemical or physical characteristics, termed "difficulty", that make food items either low in value or unpalatable to most herbivores, and (4) relevant temporal and spatial scales at which diets and niche breadth were measured. Obligatory specialists always have a narrow realized niche consisting of difficult food items, and morphological adaptations and/or the loss of redundant behavioral flexibility that effectively limit their fundamental niches, precluding them from expanding their diet under changed environmental conditions. Facultative specialists have a consistently narrow realized niche for difficult foods during at least one spatial or temporal scale, but have a broad enough fundamental niche to allow them to expand their diet to include less difficult foods when environmental conditions allow. Facultative generalists have the broadest fundamental niche, allowing them to consume a wide variety of foods. However, they may occasionally demonstrate a narrow realized niche, focused on less difficult plants than is the case with specialists. Finally, the obligatory generalists always have a wide realized niche because of a relatively narrow fundamental niche, precluding them from eating much of any difficult plant. We summarize hypothesized characteristics of mammalian herbivores in each of the four categories of specialization. We demonstrate the need for further work on defining the realized and fundamental niches, comparing among herbivores across categories conducted under similar conditions, and understanding the nature of trade-offs required for specialization and generalization for both community and phylogenetically based analyses.

Type II functional responses are frequently observed in herbivores feeding in patches where plants are concentrated in space. We tested a mechanistic model of regulation of intake rate of herbivores foraging in food—concentration patches (Laca and Demment 1992, Spalinger and Hobbs 1992) that accounts for asymptotic, Type II responses. The model is based on the hypothesis that competition between cropping and chewing regulates instataneous intake rate in response to changes in the size of bites obtained by the forager. We tested this hypothesis and examined the ability of our model to account for observations of intake rate of 12 species of mammalian herbivores ranging in body mass over 4 orders of magnitude. We measured short—term intake rates of mammalian herbivores feeding in hand—assembled patches of plants. We varied bite size by changing plant height and density in patches offered to herbivores, and observed dry matter intake rates in response to this variation. Averaged across species, our model acc...

There appear to be two dominant forms of the functional response of vertebrate herbivores, corresponding to animals foraging in spatially concentrated or spatially dispersed food patches. We examine the factors contributing to the functional response of herbivores feeding on spatially concentrated browses. Specifically, we tested the hypothesis that the instantaneous intake rate of four boreal browsers (moose, Alces alces, woodland caribou, Rangifer tarandus caribou, white-tailed deer, Odocoileus virginianus, and snowshoe hares, Lepus americanus) feeding on red maple (Acer rubrum) browse is an asymptotic function of the size of bite obtainable by the herbivore. Based on this functional response model, we also hypothesized that (i) cropping rate of the herbivores would decline curvilinearly with bite size, (ii) intake rate would decline with increasing fibrousness (measured, as the neutral detergent fiber of the foods consumed), and (iii) maximum intake rate would scale as the 2/3 power of herbivore body mass (BM). Intake rates of all animals increased 2- to 10-fold with increasing bite size of stems and leaves, and conformed to the hypothesized asymptotic relation. Similarly, cropping rate declined curvilinearly with increasing bite size, and intake rate declined with increasing fibrousness of foods consumed. Maximum intake rate scaled with BM0.76, not significantly different from the hypothesized exponent of 0.67. These data support the hypothesis that bite size affects intake rate by controlling the ratio of cropping and chewing, and that the asymptotic intake rate of herbivores is related to the food processing capability of the mouth.

Plant secondary metabolites (PSMs) strongly influence diet selection by mammalian herbivores. Concentrations of PSMs vary within and among plant species, and across landscapes. Therefore, local adaptations may cause different populations of herbivores to differ in their ability to tolerate PSMs. Here, we tested the food intake responses of three populations of a marsupial folivore, the common brushtail possum (Trichosurus vulpecula Kerr), from different latitudes and habitat types, to four types of PSMs. We found clear variation in the responses of northern and southern Australian possums to PSMs. Brushtail possums from southern Australia showed marked decreases in food intake in response to all four PSMs, while the two populations from northern Australia were not as sensitive and their responses did not differ from one another. These results were unexpected, based on our understanding of the experiences of these populations with PSMs in the wild. Our results suggest that geographically separated populations of possums may have evolved differing abilities to cope with PSMs, as a result of local adaptation to their natural environments. Our results provide the basis for future studies to investigate the mechanisms by which populations of mammalian species differ in their ability to tolerate PSMs.

Sagebrush (Artemisia tridentata) comprises up to 99% of the winter and 50% of the summer diets of pygmy rabbits (Brachylagus idahoensis). Few animals specialize on such plants as sagebrush, which contain high levels of plant chemicals that can be toxic. We investigated the nutritional requirements of pygmy rabbits and their ability and propensity to consume sagebrush alone and as part of a mixed diet. We compared diet choices of pygmy rabbits with that of a generalist forager, the eastern cottontail (Sylvilagus floridanus). Pygmy rabbits had a moderately low nitrogen requirement (306.5 mg N/kg(0.75)/d), but a relatively high energy requirement, needing 750.8 kJ digestible energy/kg(0.75)/d to maintain their body mass while residing in small cages. They digested fiber in pelleted diets similarly to other small hindgut fermenters, but both cottontails and pygmy rabbits digested the fiber in sagebrush better than expected based on its indigestible acid detergent lignin content. Pygmy rabbits consumed more sagebrush than cottontails, regardless of the amount and nutritional quality of supplemental pellets provided. When consuming sagebrush alone, they ate barely enough to meet their energy requirements, whereas cottontails ate only enough sagebrush to meet 67% of theirs. Both rabbit species ate more sagebrush as the quality and quantity of supplemental pellets declined, and more greenhouse-grown sagebrush than sagebrush grown outside. Urine was more acidic when consuming sagebrush than when consuming pellets, indicating detoxification by the liver. Pygmy rabbits do not require sagebrush to survive, but seem to tolerate terpenes and other plant chemicals in sagebrush better than cottontails do.

Ultraviolet-B (UV-B) radiation has been increasing in temperate latitudes in recent decades and is expected to continue rising for some time. Enhanced UV-B radiation can change plant chemistry, yet the effects of these changes on mammalian herbivores are unknown. To examine the influence of enhanced UV-B radiation on nutrition of a specialist and generalist hindgut fermenter, we measured nutritional and chemical constituents of three common North American range plants, big sagebrush (Artemisia tridentata), yarrow (Achillea millefolium), and bluebunch wheatgrass (Pseudoregneria spicata), and how these changes influenced in vitro dry matter digestibility and in vivo digestibility by pygmy rabbits (Brachylagus idahoensis) and eastern cottontails (Sylvilagus floridanus). Forages were irradiated for 3 mo with ambient (1×) or supplemental (1.6×) UV-B radiation representing a 15% ozone depletion for Pullman, WA, USA. Enhanced UV-B radiation had minimal effects on the nutritional content and the tannin-binding capacity of forages. Similarly, the terpene concentration in sagebrush and yarrow was not affected by higher UV-B irradiances. Flavonoid compounds increased in sagebrush but decreased in yarrow. Rabbit preference and intake was not affected by treatment levels for any forage species and no differences were found between treatments for dry matter, fiber, protein digestibility, and apparent digestible energy.

Stratospheric ozone depletion has caused ground-level ultraviolet-B (UV-B) radiation to rise in temperate latitudes of both hemispheres. Because the effects of enhanced UV-B radiation on the nutrition of food consumed by mammalian herbivores are unknown, we measured nutritional and chemical constituents of 18 forages and related changes to in vitro dry matter digestibility. We also measured intake and in vivo digestibility of Pacific willow (Salix lasiandra) and alfalfa (Medicago sativa L.) by blue duikers (Cephalophus monticola). Forages were irradiated for 3 months with ambient (1x) or supplemental (1.6 x) UV-B radiation representing a 15% ozone depletion for Pullman, Washington, USA. Enhanced UV-B radiation had minimal and inconsistent effects on the nutritional content, in vitro dry matter digestibility, and protein-binding capacity of forages. However, flavonoid compounds increased in seven of the 13 forbs and woody dicots that were evaluated. Flavonoids were found to decrease only in yarrow (Achillea millefolium). When offered simultaneously, blue duikers preferred 1x and 1.6 x UV-B irradiated plants of alfalfa equally, but ate 26% less willow grown under 1.6 x UV-B radiation. However, when fed to duikers in separate feeding experiments, total dry matter intake and in vivo digestibility of dry matter, fiber, protein, and apparent energy did not differ between alfalfa and willow grown under 1x and 1.6 x UV-B radiation. We conclude that expected increases in UV-B radiation from ozone depletion would have minimal effects on intake and digestion of ruminant herbivores.

Herbivores represent the interface between primary production and higher trophic levels. The effects of enhanced UV-B radiation on microbes, invertebrate herbivores, and detritivores has received limited study in both terrestrial and aquatic ecosystems. However, although direct effects (e.g. melanoma, cataracts) on mammals have been documented, indirect effects (e.g., resulting from changes in plant chemistry) of enhanced UV-B on mammalian herbivores have not been evaluated. Although the diet of mammalian herbivores has little effect on nutritional quality for their associated predators, to the extent changes in plant chemistry affect aspects of population dynamics (e.g., growth, fecundity, densities), higher trophic levels can be affected. In this study, different forage species of varying inherent levels of key secondary metabolites are being grown in the field under either ambient or ambient plus supplemental UV-B radiation simulating a 15% stratospheric ozone depletion for Pullman, Washington. At various time intervals, foliage is being sampled and analyzed for changes in secondary metabolites and other attributes. Using controlled feeding trials, changes in plant secondary metabolites are being related to preference and digestibility in specialist and generalist mammalian hindgut herbivores, digestion in ruminants and non-ruminants, and to selected aspects of population dynamics in mammalian herbivores. Results suggest how UV-B-induced changes in plant secondary chemistry affect animal nutrition, and thus animal productivity in a range of mammalian herbivores. Reductions in palatability and digestibility of plant material along with reductions in fecundity and other aspects of population dynamics could have significant economic ramifications for farmers, ranchers and wildlife biologists.

Context Measures of intake and digestibility from captive feeding experiments are often used to evaluate the nutritional value of plant species to herbivores; however, there is question about how well plant-quality rankings from these trials predict foraging patterns of free-ranging animals. Studies addressing the alignment of results from feeding trials and herbivory in the field using captive and free-roaming conspecifics are needed. Aims Our goal was to compare the feeding patterns of snowshoe hares in captive intake and digestion trials with those of free-living conspecifics in the species’ south-western range. Methods We conducted in vivo intake and digestion trials using captive hares to determine quality and consumption levels of the predominant conifer species in our study system. In the field, we quantified browsing intensity and over-winter depletion patterns of these conifers. We then compared voluntary intake and nutritional quality measured in captivity to consumption in the field. Key results Digestible energy (DE, kJ g–1) of conifers ranged from 11.0 (Pinus contorta) to 13.8 (Pseudotsuga menziesii) among six conifers, and digestible protein (DP, g protein per 100 g feed) from 1.2 (Thuja plicata) to 2.7 (P. contorta). During digestion trials, single-species intake was correlated with the content of digestible protein (DP) and digestible energy (DE). Hares maintained energy balance when fed two single-species diets (Pinus contorta, Pseudotsuga menziesii) and a mixed-species diet. Conifer species on which hares were able to maintain body mass (Pinus contorta, Picea engelmannii, Pseudotsuga menziesii) also tended to be the most heavily exploited by free-living hares. DP content of browse species predicted both browsing intensity and overwinter depletion of conifer species. Conclusions Voluntary intake and nutritional quality of browse, especially DP, successfully predicted foraging patterns of free-ranging conspecifics. Implications Intake and digestion trials can be a useful tool for better understanding patterns of herbivory in the field, and winter habitat quality for populations in this region is likely to be influenced by access to the most energy- and protein-rich conifers.

Structural anti-quality characteristics are physical plant traits that reduce the performance and productivity of herbivores and quality of their agricultural products. Most structural anti-quality characteristics of plants affect the rate at which herbivores gather and ingest forages, reducing the total amount of food obtained or increasing the time necessary to obtain food. Structural anti-quality can substantially influence searching time (e.g., plant crypticity, distribution), cropping time (e.g., plant fibrousness, tensile and shear strength), and bite size (e.g., plant canopy structure, spinescence). Plant structural characteristics can also reduce digestion (e.g., silica), cause injury (e.g., spines, awns, burrs, calluses), or reduce the quality of animal products, such as wool (e.g., propagules). The effects of structural antiquality characteristics depend on the morphology of the herbivore, especially its size, the morphology of the focal plant, and their context within the habitat. Integrated grazing management plans should consider options to reduce the negative effects of structural anti-quality. Carefully selecting appropriate livestock species with previous experience, and the appropriate season of grazing can minimize anti-quality on rangelands. Because structural anti-quality may actually promote sustainability of grazing systems by preventing severe defoliation, or by providing refuges for highly desirable forages, it may not be desirable to completely counteract their effects. DOI:10.2458/azu_jrm_v54i4_laca

The architecture of woody food plants forces mammalian herbivores to make compromises in their food choices. Rapid rates of dry matter intake can be achieved by choosing large bites. For woody plants, however, such bites are low in nutritive quality relative to small bites taken from leaves or twigs near the growing point of the plant. This trade-off between food quality and food intake rate is central to diet optimization in browsing herbivores. We developed a model that predicts a quantitative solution to 'optimal bite size' (i.e., the bite that results in the greatest daily net energy intake) based on constraints in harvesting and digesting foods. This model responds to the chemistry and morphology of plants, and the size and digestive strategy (ruminant versus hindgut fermenter) of the herbivore. We tested the model by conducting a set of experiments in which we offered six species of dormant deciduous trees common to the boreal forests of Sweden to captive roe deer (Capreolus capreolus), red deer (Cervus elaphus), and moose (Alces alces). We also tested alternative hypotheses that animals crop bites merely in response to the morphological structure of twigs, or the distribution of twig sizes on trees. Twig diameters cropped by these animals were positively correlated with the diameter at current annual growth. However, structural measures of the trees alone were not sufficient to predict differences in choices of twig diameters among animal species. In contrast, the optimal bite size model accounted for the different bite sizes selected by animals of different sizes and explained 86% of the variation in twig diameter cropped for all plant and animal species. Hence, we concluded that our model is useful for predicting, a priori, the twig diameters selected by herbivores on dormant deciduous trees. We suggest possible ways to enhance the model and how it can be used to assess forage availability, potential diet quality, and the vulnerability of trees to herbivory.

1. We developed a simple model of foraging locomotion for mammalian herbivores that contains two parameters, maximum foraging velocity (V max , m s -1 ) and acceleration (a o , m s -2 ) and a single dependent variable, the distance between plants (d, m). 2. We examined predictions of our model by varying the distance between plants offered to nine mammalian herbivores ranging in mass from 0.05 to 533 kg. 3. Application of simple laws of motion were adequate to explain the dynamics of foraging locomotion. As the animals travelled between food items, they accelerated to a maximum foraging velocity, which they maintained until they approached the next food item. Acceleration was achieved by increasing both the length and the frequency of strides. Mean foraging velocity increased up to 10-fold as plant spacing increased, but as plant spacing increased, mean velocity was asymptotic with distance between plants. 4. V max scaled in proportion to M M0.04, and a o scaled as M -0.17 . 5. Our results suggest that small herbivores achieve similar maximum foraging velocities as do large herbivores, but can accelerate faster and therefore encounter plants more rapidly when plants are spaced closely together (