Meghan J. Camp

Examining functional components of cover: the relationship between concealment and visibility in shrub‐steppe habitat

The term “cover” has been used broadly in ecology, with a wide range of meanings, from thermal cover to security cover, to escape cover. Some habitat features could provide both thermal and security cover, or both concealment and escape cover; but in other cases, habitat features such as vegetation could impose a tradeoff between opposing functions of cover. Cover that conceals an animal from a predator also could reduce the animals visibility and thus, its ability to detect a predator early enough to escape capture. We quantified the opposing functional properties of cover (concealment and visibility) and evaluated the relationship between these properties using continuous measures in sagebrush-steppe and grassland habitats. We hypothesized that concealment and visibility would be inversely related and that the slope of this relationship would differ among sites with varying density and patchiness of shrub vegetation, imposing differing tradeoff scenarios. Concealment and visibility were inversely, but not perfectly related, implying that animals must make tradeoffs between the properties of cover, but they could achieve higher levels of one property while giving up relatively less of the other. In addition, we examined potential tradeoffs by pygmy rabbits (Brachylagus idahoensis) by comparing concealment and visibility at locations used by rabbits with measurements collected at random locations. At a small scale, pygmy rabbits traded off visibility for concealment in dense and sparse vegetation, but not in patchy vegetation. Although cover is an intuitively simple concept, it is functionally more complex, and this study provides insight into the opposing mechanisms of cover that might influence habitat use. Our work provides an initial step towards more fully understanding how cover functionally relates to predation risk.

Modeling Trade-Offs Between Plant Fiber and Toxins: A Framework for Quantifying Risks Perceived by Foraging Herbivores

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.

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Summary Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.

Grazing in sagebrush rangelands in western North America: implications for habitat quality for a sagebrush specialist, the pygmy rabbit

Livestock grazing is one of the primary uses of sagebrush rangelands in western North America; therefore, an understanding of the ecological implications of grazing on habitat quality for sagebrush-dependent wildlife is needed to help land managers balance multiple objectives for land use. We studied effects of cattle grazing on components of habitat for an uncommon sagebrush habitat specialist, the pygmy rabbit (Brachylagus idahoensis), which has been petitioned for endangered or threatened status in the USA. We evaluated multiple components of habitat before and after grazing in replicated control and treatment plots in a mesic, high-elevation sagebrush-steppe environment in south-western Montana, USA. We predicted that grazing would decrease the biomass of herbaceous forage, alter security cover, and increase rate of collapse of rabbit burrows, and we expected that these effects would be more pronounced during summer than spring. As expected, cattle grazing reduced the biomass of perennial grasses available to pygmy rabbits after grazing that occurred during either spring or summer, and the biomass of forbs after spring grazing. In contrast, grazing did not markedly influence the functional properties of vegetation related to predation risk or the integrity of rabbit burrow systems. In the context of the stocking rate of the allotments in our study (7.3 acres/Animal Unit Month, 2.95ha/Animal Unit Month), annual cattle grazing did not seem to markedly change habitat for pygmy rabbits in our study area; however, longer-term and higher intensity grazing might result in more pronounced habitat changes. Understanding the ecological implications of cattle grazing on habitat quality for pygmy rabbits and other sagebrush-dependent wildlife can guide conservation strategies for these species on sagebrush rangelands managed under multiple-use policies.

Understanding tradeoffs between food and predation risks in a specialist mammalian herbivore

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 risk of PSM-containing food to select lower predation risk when risks co-occurred. We compared food intake of pygmy rabbits during three double-choice trials designed to examine tradeoffs by offering animals two levels of food risk (1,8-cineole, a PSM) and predation risk (concealment cover). Rabbits ate more food at feeding stations with PSM-free food and high concealment cover. However, interactions between PSMs and cover suggested that the value of PSM-free food could be reduced if concealment is low and the value of high concealment can decrease if food contains PSMs. Furthermore, foraging decisions by individual rabbits suggested variation in tolerance of food or predation risks.

The Balancing Act of Foraging: Mammalian Herbivores Trade-Off Multiple Risks When Selecting Food Patches

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.

Seasonal temperature acclimatization in a semi-fossorial mammal and the role of burrows as thermal refuges

Small mammals in habitats with strong seasonal variation in the thermal environment often exhibit physiological and behavioral adaptations for coping with thermal extremes and reducing thermoregulatory costs. Burrows are especially important for providing thermal refuge when above-ground temperatures require high regulatory costs (e.g., water or energy) or exceed the physiological tolerances of an organism. Our objective was to explore the role of burrows as thermal refuges for a small endotherm, the pygmy rabbit (Brachylagus idahoensis), during the summer and winter by quantifying energetic costs associated with resting above and below ground. We used indirect calorimetry to determine the relationship between energy expenditure and ambient temperature over a range of temperatures that pygmy rabbits experience in their natural habitat. We also measured the temperature of above- and below-ground rest sites used by pygmy rabbits in eastern Idaho, USA, during summer and winter and estimated the seasonal thermoregulatory costs of resting in the two microsites. Although pygmy rabbits demonstrated seasonal physiological acclimatization, the burrow was an important thermal refuge, especially in winter. Thermoregulatory costs were lower inside the burrow than in above-ground rest sites for more than 50% of the winter season. In contrast, thermal heterogeneity provided by above-ground rest sites during summer reduced the role of burrows as a thermal refuge during all but the hottest periods of the afternoon. Our findings contribute to an understanding of the ecology of small mammals in seasonal environments and demonstrate the importance of burrows as thermal refuge for pygmy rabbits.

Interacting effects of ambient temperature and food quality on the foraging ecology of small mammalian herbivores

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.