Habitat and patch use by hyraxes: there’s no place like home?

Models of central place foraging predict that animals should forage more thoroughly in resource patches located closer to the central place. Travel time, cost of transporting food back to the central place, and exposure to predators should all act to increase foraging costs with increasing distance from the refuge. We examined habitat and patch use in rock hyraxes ( Procavia capensis ) inhabiting a group of kopjes in a semiarid savanna, Augrabies Falls National Park, South Africa. We tested the prediction of more intense patch use closer to the central place by measuring giving-up densities (GUDs) in experimental resource patches set at four different distances from the kopje and in two microhabitats differing in cover. Surprisingly, hyraxes had their lowest GUDs at intermediate distances from the kopje. These unexpected results suggest that the sentinel behaviour of hyraxes alters the probability of detection of predators for animals foraging away from the kopje.     

Patch use, apprehension, and vigilance behavior of Nubian Ibex under perceived risk of predation

Foraging theory predicts that animals will sacrifice feedingeffort in order to reduce predation risk. Once a forager choosesa habitat, it must decide how to allocate its foraging effort.Nubian Ibex are diurnal, social, cliff-dwelling herbivores.Many of their characteristics seem to have evolved as responsesto predation risk. In order to assess the effects that perceivedrisk of predation might have on foraging behavior of free-rangingNubian Ibex in the Negev Desert, Israel, we measured giving-updensities (GUDs) in artificial food patches and used them togauge apprehension level. (Apprehension can be defined as areduction in attention devoted to performing an activity asa consequence of reallocating attention to detecting or respondingto predation risk. A forager can also be vigilant. Vigilanceis often defined as time spent scanning the surroundings withthe head up.) We also quantified time budgeting using focalobservation of individual Nubian Ibex. Habitat preferences andpatch selectivity as a measure of apprehension were considered.In particular, we tested the effect of distance from refugeon GUDs, the effect of micropatch structure on selectivity,and the effect of distance from the refuge and group size onNubian Ibex vigilance level and apprehension. Nubian Ibex allocatetheir foraging effort more toward patches closer to the escapeterrain. At the same time, Nubian Ibex are more apprehensiveat intermediate distances from the cliff edge than nearer thecliff, and their use of vigilance increases with distance fromthe cliff edge. These results suggest that Nubian Ibex may switchfrom apprehension to a more extreme behavior of vigilance atgreater distances from the refuge. This study demonstrated theuse of antipredatory behaviors, apprehension, and vigilanceby a forager. Estimating apprehension and vigilance levels ofa forager simultaneously gives a more complete and accuratepicture of how the habitat is perceived by them and combinedwith measurements of GUD allow a more accurate assessment ofhabitat quality.     

Hazardous duty pay and the foraging cost of predation

We review the concepts and research associated with measuring fear and its consequences for foraging. When foraging, animals should and do demand hazardous duty pay. They assess a foraging cost of predation to compensate for the risk of predation or the risk of catastrophic injury. Similarly, in weighing foraging options, animals tradeoff food and safety. The foraging cost of predation can be modelled, and it can be quantitatively and qualitatively measured using risk titrations. Giving‐up densities (GUDs) in depletable food patches and the distribution of foragers across safe and risky feeding opportunities are two frequent experimental tools for titrating food and safety. A growing body of literature shows that: (i) the cost of predation can be big and comprise the forager's largest foraging cost, (ii) seemingly small changes in habitat or microhabitat characteristics can lead to large changes in the cost of predation, and (iii) a forager's cost of predation rises with risk of mortality, the forager's energy state and a decrease in its marginal value of energy. In titrating for the cost of predation, researchers have investigated spatial and temporal variation in risk, scale‐dependent variation in risk, and the role of predation risk in a forager's ecology. A risk titration from a feeding animal often provides a more accurate behavioural indicator of predation risk than direct observations of predator‐inflicted mortality. Titrating for fear responses in foragers has some well‐established applications and holds promise for novel methodologies, concepts and applications. Future directions for expanding conceptual and empirical tools include: what are the consequences of foraging costs arising from interference behaviours and other sources of catastrophic loss? Are there alternative routes by which organisms can respond to tradeoffs of food and safety? What does an animal's landscape of fear look like as a spatially explicit map, and how do various environmental factors affect it? Behavioural titrations will help to illuminate these issues and more.     

Rodent foraging is affected by indirect, but not by direct, cues of predation risk

We used foraging trays to determine whether oldfield mice, Peromyscuspolionotus, altered foraging in response to direct cues of predationrisk (urine of native and nonnative predators) and indirectcues of predation risk (foraging microhabitat, precipitation,and moon illumination). The proportion of seeds remaining ineach tray (a measure of the giving-up density [GUD]) was usedto measure risk perceived by mice. Mice did not alter theirGUD when presented with cues of native predators (bobcats, Lynxrufus, and red foxes, Vulpes vulpes), recently introduced predators(coyotes, Canis latrans), nonnative predators (ocelots, Leoparduspardalis), a native herbivore (white-tailed deer, Odocoileusvirginianus), or a water control. Rather, GUD was related tomicrohabitat: rodents removed more seeds from foraging trayssheltered beneath vegetative cover compared with exposed traysoutside of cover. Rodents also removed more seeds during nightswith precipitation and when moon illumination was low. Our resultssuggest that P. polionotus used indirect cues rather than directcues to assess risk of vertebrate predation. Indirect cues maybe more reliable than are direct scent cues for estimating riskfrom multiple vertebrate predators that present the most riskin open environments.     

The effects of water on patch use by two Simpson Desert granivores (Corvus coronoides and Pseudomys hermannsburgensis)

Abstract Water loss while foraging may affect the overall value of food to desert animals. When water is scarce, foragers may alter activity and shun certain types of food due to elevated water loss. When water is abundant, foragers can exploit food patches more thoroughly and remain active over a broader range of ambient conditions. In short, food and water may be complementary resources. The presence of water raises the marginal value of food, particularly those foods low in water content. We tested for the complementarity of food and water to foragers at a sand dune site in the Simpson Desert of arid Australia. To do so, we quantified patch exploitation of foragers in the presence or absence of bowls filled with water. In order to quantify patch use, we provisioned feeding trays with granulated peanuts mixed into a sand substrate. In these trays we measured giving-up densities (GUD; the amount of food left in a tray after a foraging bout) of diurnal (mostly Australian ravens, Corvus coronoides) and nocturnal foragers (mostly sandy inland mouse, Pseudomys hermannsburgensis). The presence of water affected the GUD of ravens but not of rodents. For the ravens, GUD dropped about 50% in response to added water. For ravens, water and food are strongly complementary. In addition, ravens had lower GUD in the open than the bush microhabitat, and lower GUD at the bottom than the tops of sand dunes.     

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.     

Competition between birds and mammals: A comparison of giving-up densities between crested larks and gerbils

We combined the concept of mechanisms of co-existence with the approach of giving-up densities to study inter-taxon competition between seed-eating birds and mammals. We measured feeding behaviour in food patches to define and study the guild of seed-eating vertebrates occupying sandy habitats at Bir Asluj, Negev Desert, Israel. Despite a large number of putatively granivorous rodents and birds at the site, two gerbil species (Allenbys gerbil, Gerbillus allenbyi, and the greater Egyptian gerbil, G. pyramidum) dominated nocturnal foraging, and a single bird species (crested lark, Galerida cristata) contributed all of the daytime foraging. We used giving-up densities to quantify foraging behaviour and foraging efficiencies. A low giving-up density demonstrates the ability of a forager to profitably harvest food at low abundances and to profitably utilize the foraging opportunities left behind by the less efficient forager. Gerbils had lower giving-up densities in the bush than open microhabitat, and lower giving-up densities in the semi-stabilized than stabilized sand habitats. Crested larks showed the opposite: lower giving-up densities in the open than bush, and on the stabilized than semi-stabilized sand habitats. Despite these patterns, gerbils had substantially lower giving-up densities than crested larks in both microhabitats, all sand habitats, and during each month. Several mechanisms may permit the crested lark to co-exist with the gerbils. Larks may be cream skimmers on the high spatial and temporal variability in seed abundances. Larks may rely on insects, fruit or smaller seeds. Or, larks may rely on adjacent rocky habitats.     

Constrating short-term and long-term effects of predation risk on consumer habitat use and resources

I contrasted the short-term and long-term effects of predationrisk on snail habitat use and resource dynamics. Pulmonate snails(Physella gyrina) were placed into experimental pools and exposedto four levels of predation risk while holding their densityconstant. Periphyton resources were made available in two habitats:open and covered. I hypothesized that a behavioral responseby snails to predation risk would influence periphyton standingcrop in the open and covered habitats. Snails responded to increasingpredation risk by moving into safer (covered) habitats, andthe magnitude of their response was sensitive to the actuallevel of risk: intermediate levels of risk resulted in intermediatehabitat use. However, use of the risky (open) habitat by snailswas time dependent Snails initially responded strongly to predationrisk, but they exhibited similar patterns of habitat use atall risk levels by the end of the experiment Periphyton standingcrop was positively related to predation risk. In contrast tosnail habitat use, this response was initially weak and becamestronger as the experiment progressed. Thus, the short-and long-termresponses of snail habitat use and periphyton standing cropcontrasted sharply. I suggest that the changing patterns ofsnail habitat use over time are consistent with the idea thatsnails balance predation risk against foraging gains when selectinghabitats and that the manner in which they balance foraginggains and predation risk determines the pattern of periphytonstanding crops across habitats