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Conservation havens free of invasive predators are increasingly relied upon for fauna conservation, although havened populations can lose anti-predator traits, likely making them less suitable for life 'beyond the fence'. Sustaining low levels of mammalian predator pressure inside havens may prevent the loss of anti-predator traits from havened populations. We opportunistically compared behavioural and morphological anti-predator traits between four woylie (Bettongia penicillata ogilbyi) populations- one haven isolated from all mammalian predators, one haven containing a native mammalian predator (chuditch; Dasyurus geoffroii), and their respective non-havened counterparts (each containing both chuditch and invasive predators). Havened woylies existing without mammalian predators were smaller (shorter hindfeet, smaller body weight) and less reactive (consumed more food from fox-treated and control feeding stations, less agitated during human handling) than a non-havened reference population. However, in the haven containing chuditch, we found no difference in behaviour or morphology compared to the adjacent non-havened population. Across populations, anti-predator responses tended to appear stronger at sites with higher predator activity, suggestive of an adaptive response across a gradient of predation pressure. Our findings suggest that maintaining mammalian predation pressure in conservation havens could be effective for preventing or slowing the loss of anti-predator traits from these populations.

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Habitat loss and change are often implicated as the primary causes of species extinction. Although any population can be instantly imperiled by catastrophe, most habitat loss occurs gradually, thus enabling affected individuals an adaptive advantage to occupy the best of their dwindling opportunities. I demonstrate how to infer the advantage between two habitats for any density and frequency-dependent strategy of habitat selection. I explore the concept of an Adaptive Dispersal Strategy Landscape to reveal the Evolutionarily Stable Strategy separately for ideal-free and ideal preemptive habitat selectors. Both solutions reveal an initially counterintuitive expectation that individuals living at high density gain insufficient adaptive advantage to disperse from a deteriorating habitat. Adaptive dispersal is constrained at high density because habitats of better quality are fully occupied. I test the theory with measures of movement and foraging in crossover experiments on a seminatural population of meadow voles. The experiment allowed the voles to choose among patches and between enclosures in which I differentially manipulated food and shelter. Although photographs from an infrared camera documented voles venturing from one habitat to the other, none became resident. Voles preferentially foraged in the richer of the two enclosures, even when I reversed treatments, and they foraged more in patches protected by mulched straw. The adaptive advantage of dispersal using a surrogate fitness proxy based on the voles' giving-up densities mirrored that generated by theory. The convergence between theory and experiment yields much-needed insight into our ability to test, predict, and hopefully resolve, the ecological, evolutionary, and conservation consequences of habitat loss.

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Spatial and temporal variation in perceived predation risk is an important determinant of movement and foraging activity of animals. Foraging in this landscape of fear, individuals need to decide where and when to move, and what resources to choose. Foraging theory predicts the outcome of these decisions based on energetic trade-offs, but complex interactions between perceived predation risk and preferences of foragers for certain functional traits of their resources are rarely considered. Here, we studied the interactive effects of perceived predation risk on food trait preferences and foraging behavior in bank voles (Myodes glareolus) in experimental landscapes. Individuals (n = 19) were subjected for periods of 24 h to two extreme, risk-uniform landscapes (either risky or safe), containing 25 discrete food patches, filled with seeds of four plant species in even amounts. Seeds varied in functional traits: size, nutrients, and shape. We evaluated whether and how risk modifies forager preference for functional traits. We also investigated whether perceived risk and distance from shelter affected giving-up density (GUD), time in patches, and number of patch visits. In safe landscapes, individuals increased time spent in patches, lowered GUD and visited distant patches more often compared to risky landscapes. Individuals preferred bigger seeds independent of risk, but in the safe treatment they preferred fat-rich over carb-rich seeds. Thus, higher densities of resource levels remained in risky landscapes, while in safe landscapes resource density was lower and less diverse due to selective foraging. Our results suggest that the interaction of perceived risk and dietary preference adds an additional layer to the cascading effects of a landscape of fear which affects biodiversity at resource level.

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At a global scale, organisms are under threat due to various kinds of environmental changes, such as artificial light at night (ALAN), noise, climatic change and vegetation destruction. Usually, these changes co-vary in time and space and may take effect simultaneously. Although impacts of ALAN on biological processes have been well documented, our knowledge on the combined effects of ALAN and other environmental changes on animals remains limited. In this study, we conducted field experiments in semi-natural enclosures to explore the combined effects of ALAN and vegetation height on foraging behavior, vigilance, activity patterns and body weight in dwarf striped hamsters (Cricetulus barabensis), a nocturnal rodent widely distributed in East Asia. We find that ALAN and vegetation height affected different aspects of behavior. ALAN negatively affected search speed and positively affected handling speed, while vegetation height negatively affected giving-up density and positively affected body weight. ALAN and vegetation height also additively shaped total time spent in a food patch. No significant interactive effect of ALAN and vegetation height was detected. C. barabensis exposed to ALAN and short vegetation suffered a significant loss in body weight, and possessed a much narrower temporal niche (i.e. initiated activity later but became inactive earlier) than those under other combinations of treatments. The observed behavioral responses to ALAN and changes in vegetation height may bring fitness consequences, as well as further changes in structure and functioning of local ecosystems.

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The way species use their habitat dictates their intra- and interspecific interactions. We studied the effects of the microhabitat type and slope on the movement behaviour of the Saharan horned viper (Cerastes cerastes) in its natural habitat. This viper occurs in sand dunes and moves mostly by sidewinding. Additionally, we studied the microhabitat preference of desert rodents-the vipers' main prey. We placed the vipers on different natural dune slopes and recorded their behaviour. We found a strong anti-gravitactic response: vipers moved more frequently towards the top of the dune than in any other direction, despite a decrease in stride length with increasing slope. The foraging-related behaviour of the vipers was concentrated in the dune semi-stable areas rather than its stable or shifting sand areas. We measured rodent activity by placing seed trays in the dune allowing the rodents to collect seeds. Rodent activity was the highest in the shifting sands, closely followed by the semi-stable microhabitat. These results suggest the vipers use the semi-stable microhabitat mainly for foraging and may use the shifting sand areas as commuting routes between such areas. This study may be of use for conservation efforts of psammophilic species in desert dunes.

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Finding a common currency for benefits and hazards is a major challenge in optimal foraging theory, often requiring complex computational methods. We present a new analytic approach that builds on the Marginal Value Theorem and giving-up densities while incorporating the nonlinear effect of predation risk. We map the space of all possible environments into strategy regions, each corresponding to a discrete optimal strategy. This provides a generalised quantitative measure of the trade-off between foraging rewards and hazards. This extends a classic optimal diet choice rule-of-thumb to incorporate the hazard of waiting for better resources to appear. We compare the dynamics of optimal decision-making for three foraging life-history strategies: One in which fitness accrues instantly, and two with delays before fitness benefit is accrued. Foragers with delayed-benefit strategies are more sensitive to predation risk than resource quality, as they stand to lose more fitness from a predation event than instant-accrual foragers.

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Perceived predation risk varies in space and time. Foraging in this landscape of fear alters forager-resource interactions via cascading nonconsumptive effects. Estimating these indirect effects is difficult in natural systems. Here, we applied a novel measure to quantify the diversity at giving-up density that allows to test how spatial variation in perceived predation risk modifies the diversity of multispecies resources at local and regional spatial levels. Furthermore, we evaluated whether the nonconsumptive effects on resource species diversity can be explained by the preferences of foragers for specific functional traits and by the forager species richness. We exposed rodents of a natural community to artificial food patches, each containing an initial multispecies resource community of eight species (10 items each) mixed in sand. We sampled 35 landscapes, each containing seven patches in a spatial array, to disentangle effects at local (patch) and landscape levels. We used vegetation height as a proxy for perceived predation risk. After a period of three nights, we counted how many and which resource species were left in each patch to measure giving-up density and resource diversity at the local level (alpha diversity) and the regional level (gamma diversity and beta diversity). Furthermore, we used wildlife cameras to identify foragers and assess their species richness. With increasing vegetation height, i.e., decreasing perceived predation risk, giving-up density, and local alpha and regional gamma diversity decreased, and patches became less similar within a landscape (beta diversity increased). Foragers consumed more of the bigger and most caloric resources. The higher the forager species richness, the lower the giving-up density, and alpha and gamma diversity. Overall, spatial variation of perceived predation risk of foragers had measurable cascading effects on local and regional resource species biodiversity, independent of the forager species. Thus, nonconsumptive predation effects modify forager-resource interactions and might act as an equalizing mechanism for species coexistence.

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An important but understudied modality for eavesdropping between predators and prey is olfaction, especially between non-mammalian vertebrate predators and their prey. Here we test three olfactory eavesdropping predictions involving an apex reptilian predator, the sand goanna Varanus gouldii, and several species of its small mammalian prey in arid central Australia: 1) small mammals will recognize and avoid the odour of V. gouldii; 2) V. gouldii will be attracted to the odour of small mammals, especially of species that maximize its energetic returns; and 3) small mammals will be less mobile and will show higher burrow fidelity where V. gouldii is absent compared with where it is present. As expected, we found that small mammals recognized and avoided faecal odour of this goanna, feeding less intensively at food patches where the odour of V. gouldii was present than at patches with no odour or a pungency control odour. Varanus gouldii also was attracted to the odour of small mammals in artificial burrows and dug more frequently at burrows containing the odour of species that were energetically profitable than at those of species likely to yield diminishing returns. Our third prediction received mixed support. Rates of movement of three species of small mammals were no different where V. gouldii was present or absent, but burrow fidelity in two of these species increased as expected where V. gouldii had been removed. We conclude that olfaction plays a key role in the dynamic interaction between V. gouldii and its mammalian prey, with the interactants using olfaction to balance their respective costs of foraging and reducing predation risk. We speculate that the risk of predation from this apex reptilian predator drives the highly unusual burrow-shifting behaviour that characterizes many of Australia's small desert mammals.

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Foraging by consumers acts as a biotic filtering mechanism for biodiversity at the trophic level of resources. Variation in foraging behaviour has cascading effects on abundance, diversity, and functional trait composition of the community of resource species. Here we propose diversity at giving-up density (DivGUD), i.e. when foragers quit exploiting a patch, as a novel concept and simple measure quantifying cascading effects at multiple spatial scales. In experimental landscapes with an assemblage of plant seeds, patch residency of wild rodents decreased local α-DivGUD (via elevated mortality of species with large seeds) and regional γ-DivGUD, while dissimilarity among patches in a landscape (ß-DivGUD) increased. By linking theories of adaptive foraging behaviour with community ecology, DivGUD allows to investigate cascading indirect predation effects, e.g. the ecology-of-fear framework, feedbacks between functional trait composition of resource species and consumer communities, and effects of inter-individual differences among foragers on the biodiversity of resource communities.

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ABSTRACT Cities have grown throughout the Andes and we know little about the ecology of those species that tolerate them, limiting our options to do conservation. We applied optimal foraging theory to examine the behavior of the Andean White-eared Opossum (Didelphispernigra), in a suburban area in Bogotá, Colombia. We used the giving-up density technique, which uses the amount of food left in a feeding patch, to evaluate whether the opossum's foraging costs were affected by the height of food from the ground, and the quality and quantity of food. We also evaluated whether the spatial heterogeneity of the study site affected the opossum's foraging. We used an artificial feeding patch to test these ideas. When food was either concentrated and, in less amount (concentrated food) or diluted and more amount (diluted food), the opossums preferred to forage at 2 m than at 0.5 m, and they showed a preference for concentrated food at 0.5 m, but not at 2 m. The opossums' habitat use was affected by the spatial heterogeneity at the study site and animals preferred foraging along metal fences than on live fences made of trees. When a cable allowed connection between the metallic and live fences, the value of food patches at the live fence appeared to increase. Thus, although the opossums need resources associated with natural environments, our results suggest that there are human modifications that can benefit them, such as those that reduce the risk of predation and favor their mobility in suburban environments.

RESUMEN En los Andes las ciudades han crecido y sabemos poco sobre la ecología de aquellas especies que toleran dicho crecimiento, limitando las opciones para su apropiada conservación. Aplicamos la teoría de uso óptimo de parches para examinar el comportamiento de forrajeo del fara (Didelphispernigra) en un área suburbana de Bogotá, Colombia. Utilizamos la técnica de densidad de abandono, que usa la cantidad de alimento dejada en un parche, para evaluar si los costos de forrajeo de este marsupial fueron afectados por la altura de los parches desde el suelo, así como la calidad y cantidad de alimento. Igualmente, evaluamos el efecto de la heterogeneidad espacial del área de estudio sobre el forrajeo del fara. Usamos un parche alimentario artificial con alimento líquido para evaluar estas hipótesis. Cuando el alimento era concentrado y en menor cantidad (concentrado) o diluido y en mayor cantidad (diluido), los faras prefirieron forrajear a 2 m, y prefirieron el alimento concentrado sobre el diluido a 0.5 m, pero no a 2 m. El uso del hábitat del fara fue afectado por la heterogeneidad espacial en el sitio de estudio y ellos prefirieron forrajear a lo largo de cercas metálicas, más que sobre cercas vivas. Cuando un cable permitió la conexión entre las cercas metálicas y vivas, el valor de algunos parches en la cerca viva pareció aumentar. Aunque los faras necesitan recursos asociados con ambientes naturales, nuestros resultados sugieren que las modificaciones humanas, como las que reducen el riesgo de depredación y favorecen su movilidad, pueden beneficiarlos.

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Foraging is risky and involves balancing the benefits of resource acquisition with costs of predation. Optimal foraging theory predicts where, when and how long to forage in a given spatiotemporal distribution of risks and resources. However, significant variation in foraging behaviour and resource exploitation remain unexplained. Using single foragers in artificial landscapes of perceived risks and resources with diminishing returns, we aimed to test whether foraging behaviour and resource exploitation are adjusted to risk level, vary with risk during different components of foraging, and (co)vary among individuals. We quantified foraging behaviour and resource exploitation for 21 common voles (Microtus arvalis). By manipulating ground cover, we created simple landscapes of two food patches varying in perceived risk during feeding in a patch and/or while travelling between patches. Foraging of individuals was variable and adjusted to risk level and type. High risk during feeding reduced feeding duration and food consumption more strongly than risk while travelling. Risk during travelling modified the risk effects of feeding for changes between patches and resulting evenness of resource exploitation. Across risk conditions individuals differed consistently in when and how long they exploited resources and exposed themselves to risk. These among-individual differences in foraging behaviour were associated with consistent patterns of resource exploitation. Thus, different strategies in foraging-under-risk ultimately lead to unequal payoffs and might affect lower trophic levels in food webs. Inter-individual differences in foraging behaviour, i.e. foraging personalities, are an integral part of foraging behaviour and need to be fully integrated into optimal foraging theory.

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The marginal value theorem is an optimal foraging model that predicts how efficient foragers should respond to both their ecological and social environments when foraging in food patches, and it has strongly influenced hypotheses for primate behavior. Nevertheless, experimental tests of the marginal value theorem have been rare in primates and observational studies have provided conflicting support. As a step towards filling this gap, we test whether the foraging decisions of captive chimpanzees (Pan troglodytes) adhere to the assumptions and qualitative predictions of the marginal value theorem. We presented 12 adult chimpanzees with a two-patch foraging environment consisting of both low-quality (i.e., low-food density) and high-quality (i.e., high-food density) patches and examined the effect of patch quality on their search behavior, foraging duration, marginal capture rate, and its proxy measures: giving-up density and giving-up time. Chimpanzees foraged longer in high-quality patches, as predicted. In contrast to predictions, they did not depress high-quality patches as thoroughly as low-quality patches. Furthermore, since chimpanzees searched in a manner that fell between systematic and random, their intake rates did not decline at a steady rate over time, especially in high-quality patches, violating an assumption of the marginal value theorem. Our study provides evidence that chimpanzees are sensitive to their rate of energy intake and that their foraging durations correlate with patch quality, supporting many assumptions underlying primate foraging and social behavior. However, our results question whether the marginal value theorem is a constructive model of chimpanzee foraging behavior, and we suggest a Bayesian foraging framework (i.e., combining past foraging experiences with current patch sampling information) as a potential alternative. More work is needed to build an understanding of the proximate mechanisms underlying primate foraging decisions, especially in more complex socioecological environments.

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Artificial light at night has greatly changed the physical environment for many organisms on a global scale. As an energy efficient light resource, light emitting diodes (LEDs) have been widely used in recent years. As LEDs often have a broad spectrum, many biological processes may be potentially affected. In this study, we conducted manipulated experiments in rat-proof enclosures to explore the effects of LED night lighting on behavior of a nocturnal rodent, the Mongolian five-toed jerboa (Allactaga sibirica). We adopted the giving-up density (GUD) method and camera video trapping to study behavioral responses in terms of patch use, searching efficiency and vigilance. With the presence of white LED lighting, jerboas spent less time in patches, foraged less intensively (with higher GUDs) and became vigilant more frequently, while their searching efficiency was higher than under dark treatment. Although both positive and negative effects of LEDs on foraging were detected, the net effect of LEDs on jerboas is negative, which may further translate into changes in population dynamics, inter-specific interaction and community structure. To our knowledge, this is the first field study to explore how LED lighting affect foraging behavior and searching efficiency in rodents. Our results may have potential implications for practices such as pest control.

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An optimal habitat-selecting organism should use a dispersal strategy that enables occupation of the habitat yielding greatest fitness. The strategy is complicated when habitat quality varies through time. Theory predicts that the long-term distribution of individuals will match mean habitat quality while undermatching current habitat quality. I tested the prediction with experiments on controlled populations of meadow voles occupying two pairs of field enclosures. I released equal numbers, and equal sexes, of voles in each enclosure, and varied resource abundance between enclosures by supplemental feeding. I measured the voles' response with giving-up densities (GUDs) in artificial foraging patches, and with live-trapping at the end of the experiment. The data were consistent with only one of four a priori dispersal models. Giving-up densities declined with resource supply because short-term supply had no effect on population density. GUDs were invariant to the time course of the experiment because densities were proportional to each enclosure's long-term mean quality. Similar patterns in sex ratios and patterns of habitat occupation by juvenile voles born during the experiment reinforce the interpretation of time-averaged habitat matching. This study adds to the cumulating evidence that strategies of space use converge toward behavioral and evolutionary optima.

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How rodents perceive predation risk may alter their seed foraging behaviour and therefore potentially influence the recruitment of tree species. In this study we used two methods to investigate the effect of predation risk on habitat use by the African giant pouched rat (Cricetomys sp. nov) in Ngel Nyaki forest reserve, Nigeria. The first method was 'giving up density' (GUD), an index of perceived risk of predation at an artificial food patch, and the second was the 'spool-and-line' approach, whereby unravelling spools attached to rodent bodies are used to trace their tracks. For our GUD experiment, we chose four major sites in the forest; two representative of core habitat and two at the forest edge. Additionally, three characteristic microsites were used in the GUD experiment: dense understory, open understory and near-burrows. We hypothesised that GUDs would be lower on every succeeding observation day as rats learn to use the food patches, higher GUDs would be observed in the forest edges and open microsites, and rats would show preference for the microhabitats with least exposure to potential predators. In support of our first hypothesis, we found that GUDs were highest on the first experimental nights of every session. We also found that GUDs in the forest edges were higher than GUDs in the forest core. Lower GUDs were observed close to the rat burrows and in dense understory microsites, even though these differences were not statistically significant. Tracking of rat movements using the spool-and-line method overall revealed an even use across microhabitats, with a weak preference for those with logs, dense understory or exposed ground. Overall, our results suggest that vegetation density on a microhabitat scale has little or no effect on the perception of predation risk by African giant pouched rats.

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Individual size is a major determinant of mobile organisms' ecology and behavior. This study aims to explore whether allometric scaling principles can provide an underlying framework for general patterns of resource patch use. To this end, we used giving-up densities (GUDs), that is, the amount of resources remaining in a patch after a forager has quit feeding, as a comparative measure of the amount of resources exploited by a forager of any given size. We specifically tested the hypothesis that size-dependent responses to both internal (energy requirement) and external (risk management) forces may have an effect on GUDs. We addressed this topic by conducting an extensive meta-analysis of published data on granivorous rodents, including 292 GUD measurements reported in 25 papers. The data set includes data on 22 granivorous rodent species belonging to three taxonomic suborders (Castorimorpha, Myomorpha, and Sciuromorpha) and spans three habitat types (desert, grassland, and forest). The observations refer to both patches subject to predation risk and safe patches. Pooling all data, we observed positive allometric scaling of GUDs with average forager size (scaling exponent = 0.45), which explained 15% of overall variance in individual GUDs. Perceived predation risk during foraging led to an increase in GUDs independently of forager size and taxonomy and of habitat type, which explained an additional 12% of overall GUD variance. The size scaling exponent of GUDs is positive across habitat types and taxonomic suborders of rodents. Some variation was observed, however. The scaling coefficients in grassland and forest habitat types were significantly higher than in the desert habitat type. In addition, Sciuromorpha and Myomorpha exhibited a more pronounced size scaling of GUDs than Castorimorpha. This suggests that different adaptive behaviors may be used in different contexts and/or from different foragers. With body size being a fundamental ecological descriptor, research into size scaling of GUDs may help to place patch-use observations in a broader allometric framework.

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ABSTRACT: Prey strategically respond to the risk of predation by varying their behavior while balancing the tradeoffs of food and safety. We present here an experiment that tests the way the same indirect cues of predation risk are interpreted by bank voles, Myodes glareolus, as the game changes through exposure to a caged weasel. Using optimal patch use, we asked wild-caught voles to rank the risk they perceived. We measured their response to olfactory cues in the form of weasel bedding, a sham control in the form of rabbit bedding, and an odor-free control. We repeated the interviews in a chronological order to test the change in response, i.e., the changes in the value of the information. We found that the voles did not differentiate strongly between treatments pre-exposure to the weasel. During the exposure, vole foraging activity was reduced in all treatments, but proportionally increased in the vicinity to the rabbit odor. Post-exposure, the voles focused their foraging in the control, while the value of exposure to the predator explained the majority of variation in response. Our data also suggested a sex bias in interpretation of the cues. Given how the foragers changed their interpretation of the same cues based on external information, we suggest that applying predator olfactory cues as a simulation of predation risk needs further testing. For instance, what are the possible effective compounds and how they change "fear" response over time. The major conclusion is that however effective olfactory cues may be, the presence of live predators overwhelmingly affects the information voles gained from these cues. SIGNIFICANCE STATEMENT: In ecology, "fear" is the strategic response to cues of risk an animal senses in its environment. The cues suggesting the existence of a predator in the vicinity are weighed by an individual against the probability of encounter with the predator and the perceived lethality of an encounter with the predator. The best documented such response is variation in foraging tenacity as measured by a giving-up density. In this paper, we show that an olfactory predator cue and the smell of an interspecific competitor result in different responses based on experience with a live-caged predator. This work provides a cautionary example of the risk in making assumptions regarding olfactory cues devoid of environmental context.

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Foraging is a three-stage process during which animals visit patches, consume food and quit. Foraging theory exploring relative patch quality has mostly focused on patch use and quitting decisions, ignoring the first crucial step for any forager: finding food. Yet, the decision to visit a patch is just as important as the decision to quit, as quitting theories can only be used if animals visit patches in the first place. Therefore, to better understand the foraging process and predict its outcomes, it is necessary to explore its three stages together. We used the common brushtail possum (Trichosurus vulpecula) as a model to investigate foraging decisions in response to food varying in quality. In particular, we tested whether patch nutritional quality affected the following: (1) patch visits; (2) behaviours at the patch during a foraging visit; and (3) patch quitting decisions (quantified using giving up density-GUD). Free-ranging possums were presented with diets varying in nitrogen content and concomitantly volatile organic compound (VOC) composition at feeding stations in the wild. We found that possums were able to distinguish between different quality foods from afar, despite the location of the diets changed daily. Possums used VOC (i.e. odour cues) emitted by the diets to find and select patches from a distance. High-quality diets with higher protein and lower fibre were visited more often and for longer. Possums spent more time foraging on diets high in nutritional content, resulting in lower GUDs. Our study provides important quantitative evidence that foraging efficiency plays out during all the three stages of the foraging process (i.e. visit, consume and quit), and demonstrates the significance of considering all these stages together in future studies and foraging models. Sensory cues such as food odours play a critical role in helping foragers, including mammalian herbivores, find high-quality food. This allows foragers to make quick, accurate and important decisions about food patches well before patch quitting decisions come into play.

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An organism's foraging strategy depends on its ability to acquire and use information from food patches. A forager that does not obtain information from a food patch should remain there for a fixed time or a fixed amount of harvest. Foragers that obtain partial or complete information from the patch should be able to estimate costs and benefits from its exploitation. The foraging strategy used by a forager can be determined using the giving-up density (GUD) technique. The GUD is the amount of food left by a forager after feeding in a patch with diminishing returns. We constructed an artificial feeder to examine the foraging strategy of the short-tailed fruit bat Carollia perspicillata (Chiroptera: Phyllostomidae). In this feeder, the bat's harvest rate declined with time, i.e., they experienced diminishing returns. When offered feeders with different initial food densities, bats consumed proportionately more from rich patches than from poor ones, but they equalized GUDs. When offered feeders with different concentrations of sugar (i.e., different energy content), the higher the sugar concentration in the food, the more they consumed. Altogether, the evidence indicates that these fruit bats used complete, unbiased information to decide how much time to stay in a food patch.

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Where direct killing is rare and niche overlap low, sympatric carnivores may appear to coexist without conflict. Interference interactions, harassment and injury from larger carnivores may still pose a risk to smaller mesopredators. Foraging theory suggests that animals should adjust their behaviour accordingly to optimise foraging efficiency and overall fitness, trading off harvest rate with costs to fitness. The foraging behaviour of red foxes, Vulpes vulpes, was studied with automated cameras and a repeated measures giving-up density (GUD) experiment where olfactory risk cues were manipulated. In Plitvice Lakes National Park, Croatia, red foxes increased GUDs by 34% and quitting harvest rates by 29% in response to wolf urine. In addition to leaving more food behind, foxes also responded to wolf urine by spending less time visiting food patches each day and altering their behaviour in order to compensate for the increased risk when foraging from patches. Thus, red foxes utilised olfaction to assess risk and experienced foraging costs due to the presence of a cue from gray wolves, Canis lupus. This study identifies behavioural mechanisms which may enable competing predators to coexist, and highlights the potential for additional ecosystem service pathways arising from the behaviour of large carnivores. Given the vulnerability of large carnivores to anthropogenic disturbance, a growing human population and intensifying resource consumption, it becomes increasingly important to understand ecological processes so that land can be managed appropriately.

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