The influence of bite size on foraging at larger spatial and temporal scales by mammalian herbivores

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.     

Olfactory and visual plant cues as drivers of selective herbivory

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

Soil nutrient status determines how elephant utilize trees and shape environments

1. Elucidation of the mechanism determining the spatial scale of patch selection by herbivores has been complicated by the way in which resource availability at a specific scale is measured and by vigilance behaviour of the herbivores themselves. To reduce these complications, we studied patch selection by an animal with negligible predation risk, the African elephant. 2. We introduce the concept of nutrient load as the product of patch size, number of patches and local patch nutrient concentration. Nutrient load provides a novel spatially explicit expression of the total available nutrients a herbivore can select from. 3. We hypothesized that elephant would select nutrient-rich patches, based on the nutrient load per 2500 m(2) down to the individual plant scale, and that this selection will depend on the nitrogen and phosphorous contents of plants. 4. We predicted that elephant would cause more adverse impact to trees of lower value to them in order to reach plant parts with higher nutrient concentrations such as bark and root. However, elephant should maintain nutrient-rich trees by inducing coppicing of trees through re-utilization of leaves. 5. Elephant patch selection was measured in a homogenous tree species stand by manipulating the spatial distribution of soil nutrients in a large field experiment using NPK fertilizer. 6. Elephant were able to select nutrient-rich patches and utilized Colophospermum mopane trees inside these patches more than outside, at scales ranging from 2500 down to 100 m(2) . 7. Although both nitrogen and phosphorus contents of leaves from C. mopane trees were higher in fertilized and selected patches, patch choice correlated most strongly with nitrogen content. As predicted, stripping of leaves occurred more in nutrient-rich patches, while adverse impact such as uprooting of trees occurred more in nutrient-poor areas. 8. Our results emphasize the necessity of including scale-dependent selectivity in foraging studies and how elephant foraging behaviour can be used as indicators of change in the availability of nutrients.     

Cattle select African savanna termite mound patches less when sharing habitat with wild herbivores

African savanna termite mounds function as nutrient‐rich foraging hotspots for different herbivore species, but little is known about their effects on the interaction between domestic and wild herbivores. Understanding such effects is important for better management of these herbivore guilds in landscapes where they share habitats. Working in a central Kenyan savanna ecosystem, we compared selection of termite mound patches by cattle between areas cattle accessed exclusively and areas they shared with wild herbivores. Termite mound selection index was significantly lower in the shared areas than in areas cattle accessed exclusively. Furthermore, cattle used termite mounds in proportion to their availability when they were the only herbivores present, but used them less than their availability when they shared foraging areas with wild herbivores. These patterns were associated with reduced herbage cover on termite mounds in the shared foraging areas, partly indicating that cattle and wild herbivores compete for termite mound forage. However, reduced selection of termite mound patches was also reinforced by higher leafiness of Brachiaria lachnantha (the principal cattle diet forage species) off termite mounds in shared than in unshared areas. Taken together, these findings suggest that during wet periods, cattle can overcome competition for termite mounds by taking advantage of wildlife‐mediated increased forage leafiness in the matrix surrounding termite mounds. However, this advantage is likely to dissipate during dry periods when forage conditions deteriorate across the landscape and the importance of termite mounds as nutrient hotspots increases for both cattle and wild herbivores. Therefore, we suggest that those managing for both livestock production and wildlife conservation in such savanna landscapes should adopt grazing strategies that could lessen competition for forage on termite mounds, such as strategically decreasing stock numbers during dry periods.     

Quantifying the response of free-ranging mammalian herbivores to the interplay between plant defense and nutrient concentrations

While trying to achieve their nutritional requirements, foraging herbivores face the costs of plant defenses, such as toxins. Teasing apart the costs and benefits of various chemical constituents in plants is difficult because their chemical defenses and nutrient concentrations often co-vary. We used an approach derived from predator–prey studies to quantitatively compare the foraging response of a free-ranging mammalian herbivore, the swamp wallaby (Wallabia bicolor), through three feeding trials with artificial diets that differed in their concentrations of (1) the terpene 1,8-cineole, (2) primary constituents (including nitrogen and fiber), and (3) both the terpene and the primary constituents. Applying the giving-up density (GUD) framework, we demonstrated that the foraging cost of food patches increases with higher dietary cineole concentration and decreases with higher dietary nutrient concentration. The effect of combined differences in nutrients and cineole concentrations on GUD was interactive, and high nutrient food required more cineole to achieve the same patch value as low nutrient food. Our results indicate that swamp wallabies equate low nutrient, poorly defended food with high nutrient, highly defended food, providing two contrasting diets with similar cost–benefit outcomes. This behavior suggests that equal concentrations of chemical defenses provide nutrient-poor plants with relatively greater protection as nutrient-rich plants. Nutrient-rich plants may therefore face the exacerbated problem of being preferred by herbivores and therefore need to produce more defense compounds to achieve the same level of defense as nutrient-poor plants. Our findings help explain the difference in anti-herbivore strategy of nutrient-poor and rich plants, i.e., tolerance versus defense.     

Patch Use in Free‐Ranging Goats: Does a Large Mammalian Herbivore Forage like Other Central Place Foragers?

Classic central place foraging theory does not focus on the foraging of central place herbivores. This is especially true with regard to large mammalian herbivores. To understand the foraging dynamics of these neglected foragers, we measured giving‐up densities (GUDs) in artificial food patches. We did this at different distances away from the central point (i.e. corral) for a herd of free‐ranging domestic goats. To determine temporal changes, we conducted the study over a 3‐mo period during an extended dry season. Throughout our study, goats foraged across a gradient of food availability where forage was more available farther away from the central point. In contrast to the prediction that predation risk and/or increased travel costs were the main drivers of foraging decisions, we found that the goats increased their feeding effort (i.e. achieved lower GUDs) the farther away they moved from the central point. This suggests that either metabolic or missed opportunity costs were the main factors that influenced foraging decisions. In addition, we suggest that social foraging may have also played a role. With increases in foraging opportunities away from the central point, a herd will likely move slowly while foraging. As a result, individuals can feed intensively from patches but remain part of the group. Ironically, owing to the sustained close proximity of other group members, individuals may perceive patches farther from the central point as being safer. Temporally, the goats increased their feeding effort throughout the dry season. This suggests there was a decline in food quality and/or availability across the environment as the study progressed. Despite this increase in feeding effort, the negative relationship with distance did not change. Ultimately, our results provide key insight into how metabolic, missed opportunity and perceived predation costs influence the feeding decisions of large central place herbivores.     

西双版纳野象谷景区亚洲象取食作用对五种植物的影响

摘要：本研究在西双版纳利用样线调查法调查亚洲象的取食作用对野象谷景区常见分布的5种植物即酸苔树、象鼻藤、粗糠柴、翅果麻和黄牛木的影响,调查中记录样线上每一目标植株的基径、高度、影响类型、影响程度等信息。结果表明：1）植物受影响的数量与其植株总数之间的相关性不大（Pearson correlation, r=0.608, P=0.277）;2）亚洲象对5种植物的偏爱程度不一样,酸苔树、象鼻藤、粗糠柴和翅果麻都是大象喜爱的食物,尤其是象鼻藤（偏爱指数=1.4855）,而黄牛木不受大象喜爱（偏爱指数=0.5855）;3）亚洲象对5种植物有不同的影响方式和影响程度,酸苔树以主干折断和枝叶损失为主,受大象的取食影响最严重,象鼻藤虽然是大象最喜爱的食物,但受到的取食影响最小,粗糠柴以枝叶损失和主干折断为主,翅果麻以枝叶损失为主,同时也是唯一被亚洲象剥皮的植物,黄牛木以枝叶损失为主,被推倒的情况最普遍。树干基部直径为3-8cm的植株最受大象偏爱。亚洲象对植物的选择性以及植株自身的特性是导致上述差异的主要原因。建议后续开展长期的调查以了解亚洲象对这些植物种群数量及群落结构的影响,同时应采取有效保护措施以避免象群在当地过于集中的现象。     

Effects of within-patch characteristics on the vulnerability of a plant to herbivory

Consumption of a focal plant by herbivores depends, not only on the physical and chemical characteristics of that plant, but also on the characteristics of the neighbouring vegetation. Consumption of focal plants has been related to their own characteristics and to the quality of the neighbouring vegetation, but the two have not been combined to examine the relative importance of focal plant and neighbouring vegetation characteristics.
We conducted a series of feeding trials to examine the relative importance of focal plant and various characteristics of neighbouring vegetation to browsing of a focal plant within vegetation patches. We planted Eucalyptus nitens seedlings of high and low nutrient status amongst vegetation patches differing in palatability, abundance and height. Generalist mammalian herbivores, red-bellied pademelons ( Thylogale billardierii ), were allowed to feed in each of these patches one at a time, and seedling consumption was recorded. Results were considered in light of the attractant-decoy and apparency hypotheses, which focus on the outcome to plants, and in terms of foraging theory, which is process-focussed.
Seedling and vegetation characteristics were both important. Seedlings of high nutrient status were preferred over those of low nutrient status. The relative quality, abundance and height of neighbouring vegetation all influenced browsing of a focal plant. Seedlings were more vulnerable amongst vegetation that was of low palatability, of low abundance, or was short. Seedling and vegetation effects were additive in two of three trials.
Results were consistent with both the attractant-decoy and apparency hypotheses, and could be explained in terms of maximising foraging efficiency. They demonstrate the need to consider characteristics of both the focal plant and its neighbouring vegetation when predicting the vulnerability of the former to browsing by generalist herbivores.     

Intraspecific Strategic Responses of African Elephants to Temporal Variation in Forage Quality

ABSTRACT Mammalian herbivores adopt foraging strategies to optimize nutritional trade-offs against restrictions imposed by body size, nutritional requirements, digestive anatomy, physiology, and the forage resource they exploit. Selective or generalist feeding strategies scale with body size across species. However, within species, where constraints should be most similar, responses to limitation have rarely been examined. We used African elephants (Loxodonta Africana) to test for changes in seasonal diet quality of individuals of differing body size and sex through measurement of fecal nitrogen and phosphorus. We measured physiological stress response of these age and sex classes to seasonal change by fecal glucocorticoid metabolite levels (i.e., stress hormones). Large body size increased tolerance to lower-quality forage. Adult males and females exhibited divergent trends; females had higher diet quality than males, irrespective of body size. When limited by forage availability or quality during the dry season, diet quality declined across all body sizes, but weaned calves ingested a higher-quality diet than larger-bodied adults. On release from restriction during the wet season, weaned calf nitrogen concentrations were consistently high and stress hormone levels decreased, whereas adult female phosphorus levels were highest and less variable and stress hormone levels were unchanged. The ability to adjust forage quality is an important strategy used to ensure adequate nutritional intake according to body size limitations. Although body size is a key determining factor of dietary differences between adult elephants, foraging strategies are also driven by specific nutritional requirements, which may override the body size effects driving foraging decisions in some cases. The diversity of intraspecific response highlights ecologically segregated entities within a species and should be a concern for population management planning, particularly for threatened species. Fecal diet quality and stress hormone analysis could provide an early and sensitive indicator for monitoring age and sex class responses to resource restriction in high-density elephant populations.     

The dilemma of foraging herbivores: dealing with food and fear

For foraging herbivores, both food quality and predation risk vary across the landscape. Animals should avoid low-quality food patches in favour of high-quality ones, and seek safe patches while avoiding risky ones. Herbivores often face the foraging dilemma, however, of choosing between high-quality food in risky places or low-quality food in safe places. Here, we explore how and why the interaction between food quality and predation risk affects foraging decisions of mammalian herbivores, focusing on browsers confronting plant toxins in a landscape of fear. We draw together themes of plant–herbivore and predator–prey interactions, and the roles of animal ecophysiology, behaviour and personality. The response of herbivores to the dual costs of food and fear depends on the interplay of physiology and behaviour. We discuss detoxification physiology in dealing with plant toxins, and stress physiology associated with perceived predation risk. We argue that behaviour is the interface enabling herbivores to stay or quit food patches in response to their physiological tolerance to these risks. We hypothesise that generalist and specialist herbivores perceive the relative costs of plant defence and predation risk differently and intra-specifically, individuals with different personalities and physiologies should do so too, creating individualised landscapes of food and fear. We explore the ecological significance and emergent impacts of these individual-based foraging outcomes on populations and communities, and offer predictions that can be clearly tested. In doing so, we provide an integrated platform advancing herbivore foraging theory with food quality and predation risk at its core.     

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.     

Herbivores as architects of savannas: inducing and modifying spatial vegetation patterning

In this paper, we address the question whether and through which mechanisms herbivores can induce spatial patterning in savanna vegetation, and how the role of herbivory as a determinant of vegetation patterning changes with herbivore density and the pre-existing pattern of vegetation. We thereto developed a spatially explicit simulation model, including growth of grasses and trees, vertical zonation of browseable biomass, and spatially explicit foraging by grazers and browsers. We show that herbivores can induce vegetation patterning when two key assumptions are fulfilled. First, herbivores have to increase the attractiveness of a site while foraging so that they will revisit this site, e.g. through an increased availability or quality of forage. Second, foraging should be spatially explicit, e.g. when foraging at a site influences vegetation at larger spatial scales or when vegetation at larger spatial scales influences the selection and utilisation of a site. The interaction between these two assumptions proved to be crucial for herbivores to produce spatial vegetation patterns, but then only at low to intermediate herbivore densities. High herbivore densities result in homogenisation of vegetation. Furthermore, our model shows that the pre-existing spatial pattern in vegetation influences the process of vegetation patterning through herbivory. However, this influence decreases when the heterogeneity and dominant scale of the initial vegetation decreases. Hence, the level of adherence of the herbivores to forage in pre-existing patches increases when these pre-existing patches increase in size and when the level of vegetation heterogeneity increases. The findings presented in this paper, and critical experimentation of their ecological validity, will increase our understanding of vegetation patterning in savanna ecosystems, and the role of plant–herbivore interactions therein.     

Opportunity costs influence food selection and giving‐up density of dabbling ducks

The interaction of animals with their food can yield insights into habitat characteristics, such as perceived predation risk and relative quality. We deployed experimental foraging patches in wetlands used by migrating dabbling ducks Anas spp. in the central Illinois River Valley to estimate variation in seed removal and giving‐up density (GUD; i.e. density of food remaining in patches following abandonment) with respect to seed density, seed size, seed depth in the substrate, substrate firmness, perceived predation risk, and an energetic profitability threshold (i.e. critical food density). Seed depth and the density of naturally‐occurring seeds outside of experimental plots affected seed removal and GUD in experimental patches more than perceived predation risk, seed density, seed size or substrate firmness. The greatest seed removal and lowest GUDs in experimental patches occurred when food resources in alternative foraging locations outside of plots (i.e. opportunity costs) appeared to be near or below a critical food density (i.e. 119–181 kg ha^–1). Giving‐up densities varied substantially from a critical food density across a range of food densities in alternative foraging locations suggesting that fixed GUDs should not be used as surrogates for critical food densities in energetic carrying capacity models. Foraging and resting rates in and near experimental foraging patches did not reflect patterns of seed removal and were poor predictors of GUD and foraging habitat quality. Our results demonstrated the usefulness of GUDs as indicators of habitat quality for subsurface, benthic foragers relative to other available foraging patches and suggested that food may be limited for dabbling ducks during spring migration in some years in the midwestern USA.     

Importance of nutritional and anti-parasite strategies in the foraging decisions of horses: an experimental test

The primary goal of foraging herbivores is to maximise the net rate of intake of digestible energy (or of a limiting nutrient). However, foraging strategies of herbivores are also sensitive to other selective forces (e.g. predation, parasites), which may modify their choice of feeding patches. Horses feed in spatially complex swards, and allocate their time among patches which vary both in terms of their nutritional benefits, and the risk of parasitism. It has long been suggested that horses allocate time among patches principally in relation to the risk of parasitism, though the nutritional costs and benefits of different foraging choices must play some role too. In this study, we investigated the roles of nutritional and anti-parasite factors in foraging decisions by horses. Six naturally parasitized and six unparasitized two-year-old geldings were allowed to graze from pairs of trays (112×72 cm) with swards at two heights (tall: 52 cm, medium: 15 cm) and two levels of contamination (280 g m⁻² of faeces, no faeces) in a latin square, pair-wise design with six different choices. In nutritional terms the differences between the swards were slight: the tall sward provided larger bite sizes and higher intake rates (+36%). Feeding motivation was not high, and the faecal stimulus was strong. The horses selected uncontaminated swards when the nutritional benefits were identical, and tall swards in virtually all circumstances. Contamination of the preferred tall sward (i.e. a test of the tradeoff between anti-parasite and nutritional strategies) had little effect on its selection by the horses. Their parasite status also had little effect: both groups selected the tall sward in the tradeoff (though this was only a trend for the non-parasitised group in one analysis). We conclude that patch selection by horses in these particular conditions was driven principally by their nutritional strategy.     

The role of pre- and post- alighting detection mechanisms in the responses to patch size by specialist herbivores

Experimental data on the relationship between plant patch size and population density of herbivores within fields often deviates from predictions of the theory of island biogeography and the resource concentration hypothesis. Here we argue that basic features of foraging behaviour can explain different responses of specialist herbivores to habitat heterogeneity. In a combination of field and simulation studies, we applied basic knowledge on the foraging strategies of three specialist herbivores: the cabbage aphid (Brevicoryne brassicae), the cabbage butterfly (Pieris rapae L.) and the diamondback moth (Plutella xylostella L.), to explain differences in their responses to small scale fragmentation of their habitat. In our field study, populations of the three species responded to different sizes of host plant patches (9 plants and 100 plants) in different ways. Densities of winged cabbage aphids were independent of patch size. Egg‐densities of the cabbage butterfly were higher in small than in large patches. Densities of diamondback moth adults were higher in large patches than in small patches. When patches in a background of barley were compared with those in grass, densities of the cabbage aphid and the diamondback moth were reduced, but not cabbage butterfly densities. To explore the role of foraging behaviour of herbivores on their response to patch size, a spatially explicit individual‐based simulation framework was used. The sensory abilities of the insects to detect and respond to contact, olfactory or visual cues were varied. Species with a post‐alighting host recognition behaviour (cabbage aphid) could only use contact cues from host plants encountered after landing. In contrast, species capable with a pre‐alighting recognition behaviour, based on visual (cabbage butterfly) or olfactory (diamondback moth) cues, were able to recognise a preferred host plant whilst in flight. These three searching modalities were studied by varying the in flight detection abilities, the displacement speed and the arrestment response to host plants by individuals. Simulated patch size – density relationships were similar to those observed in the field. The importance of pre‐ and post‐ alighting detection in the responses of herbivores to spatial heterogeneity of the habitat is discussed.     

A Device to Study the Behavioral Responses of Zooplankton to Food Quality and Quantity

In order to explore the behavioral mechanisms underlying aggregation of foragers on local resource patches, it is necessary to manipulate the location, quality and quantity of food patches. This requires careful control over the conditions in the foraging arena, which may be a challenging task in the case of aquatic resource-consumer systems, like that of freshwater zooplankton feeding on suspended algal cells. We present an experimental tool designed to aid behavioral ecologists in exploring the consequences of resource characteristics for zooplankton aggregation behavior and movement decisions under conditions where the boundaries and characteristics (quantity and quality) of food patches can be standardized. The aggregation behavior of Daphnia magna and D. galeata x hyalina was tested in relation to i) the presence or absence of food or ii) food quality, where algae of high or low nutrient (phosphorus) content were offered in distinct patches. Individuals of both Daphnia species chose tubes containing food patches and D. galeata x hyalina also showed a preference towards food patches of high nutrient content. We discuss how the described equipment complements other behavioral approaches providing a useful tool to understand animal foraging decisions in environments with heterogeneous resource distributions.     

Context dependence in foraging behaviour of Achillea millefolium

Context-dependent foraging behaviour is acknowledged and well documented for a diversity of animals and conditions. The contextual determinants of plant foraging behaviour, however, are poorly understood. Plant roots encounter patchy distributions of nutrients and soil fungi. Both of these features affect root form and function, but how they interact to affect foraging behaviour is unknown. We extend the use of the marginal value theorem to make predictions about the foraging behaviour of roots, and test our predictions by manipulating soil resource distribution and inoculation by soil fungi. We measured plant movement as both distance roots travelled and time taken to grow through nutrient patches of varied quality. To do this, we grew Achillea millefolium in the centers of modified pots with a high-nutrient patch and a low-nutrient patch on either side of the plant (heterogeneous) or patch-free conditions (homogeneous). Fungal inoculation, but not resource distribution, altered the time it took roots to reach nutrient patches. When in nutrient patches, root growth decreased relative to homogeneous soils. However, this change in foraging behaviour was not contingent upon patch quality or fungal inoculation. Root system breadth was larger in homogeneous than in heterogeneous soils, until measures were influenced by pot edges. Overall, we find that root foraging behaviour is modified by resource heterogeneity but not fungal inoculation. We find support for predictions of the marginal value theorem that organisms travel faster through low-quality than through high-quality environments, with the caveat that roots respond to nutrient patches per se rather than the quality of those patches.     

Landscape-scale habitat response of African elephants shows strong selection for foraging opportunities in a human dominated ecosystem

Balancing trade-offs between foraging and risk factors is a fundamental behavior that structures the spatial distribution of species. For African elephants Loxodonta africana, human pressures from poaching and conflict are primary drivers of species decline, but little is known about how elephants structure their spatial behavior in the face of human occupancy and predation. We seek to understand how elephants balance trade-offs between resource access, human presence and human predatory risk factors (poaching and conflict killing) in an unfenced, dynamic ecosystem where elephants persist primarily outside protected areas in community rangelands. We used tracking data from 101 elephants collected between 2001 and 2016. We investigated elephant behavior in response to landcover, topography, productivity, water, human features and human predation risk using third-order resource selection functions. We extended this analysis by employing a mixed-effects multinomial regression to identify temporal shifts in habitat use, and evaluated temporal shifts in movement patterns by estimating mean squared displacement across different productivity periods. Across periods, elephants displayed strong selection for productive areas and areas near water. Temporal shifts in habitat use showed that, during the dry period, elephants were clustered around permanent water sources where humans also congregated. At the onset of the wet period, a shift occurred where elephants moved away from permanent water and from permanent settlements towards seasonal water sources and seasonal settlements. Our findings indicate that foraging and water access are important limiting factors affecting elephants that potentially restrain their spatial responses to humans at the scale of our analysis. Given that pastoralists and elephants rely on the same resources, increasing human and livestock populations enhance pressure on shared resources and space in Africa's drylands. The long-term conservation of elephants will require approaches that reduce poaching as well as landscape level planning to prevent negative impacts from increasing competition for preferred resources.     

Providing water for goats in arid landscapes: effects on feeding effort with regard to time period, herd size and secondary compounds

In arid regions, herbivores contend with a wide range of variables that influence their foraging ability. These may include plant secondary compounds (e.g. tannins and oxalates), water availability, time of day, and herd size. To determine the relative importance of these variables for goats living in a semi-desert, we measured the remaining food resources after foraging events (giving up densities-GUDs) as an index of foraging effort in artificial food patches. Time of day and the availability of drinking water had the greatest effect on GUDs. Goats achieved lower GUDs (i.e. ate more) in the afternoon than they did during the morning. We suggest this was due to missed opportunity costs of future foraging opportunities being lower in the afternoon. Ultimately, this implies that goats made short-term foraging decisions based on estimates of future feeding opportunities. When water was available, feeding effort increased and GUDs dropped almost 50%. For goats living in an arid environment, food and water are strong complementary resources. The presence of low to moderate levels of tannic or oxalic acid in food was a slight deterrent, raising the GUDs similarly. Larger herds ate more and thus obtained lower GUDs. In addition, there was an interaction of herd size and water, suggesting that in arid environments, the effect of water intensifies with population density. Our approach allowed us to quantify and rank the effects that disparate environmental factors had on the goats' foraging costs: water>time of day>herd size>plant secondary compounds. These findings ultimately demonstrate how the provision of water may increase grazing impacts by herbivores in arid landscapes.     

Partial feeding preferences and the profitability of winter-feeding sites for brent geese

The suitability of depletion theory for predicting the distribution and movements of wintering brent geese feeding in habitat patches containing foods differing qualitatively as well as quantitatively is evaluated. By monitoring both digestibility and nutrient content of potential foods throughout the season, we assess profitability of habitat patches using assimilation rates.

We argue that these geese do not conform to the predictions of an ideal free distribution because they are constrained both by nitrogen limitation and perceived mortality risks. Instead, for most of the season they exhibited partial feeding preferences by feeding on two or more types of food each day. They fed on salt marsh plants throughout the entire wintering season. In addition, from October until March they fed for part of each day on supplementary sites that were more profitable for nitrogen. In October they fed first on intertidal algae, the most profitable source of nitrogen. When this became depleted in late autumn, they moved inland to feed initially on winter wheat, where they were subject to control shooting, then onto pastures. By mid-March the pastures were no longer a significantly more profitable source of nitrogen. The geese then switched to feeding only on the salt marshes at a cost of a 39% decrease in their overall assimilation rates.

The nitrogen limitation hypothesis was supported by results of experimentally altering the nitrogen content of pasture swards. Feeding preferences correlated positively with changes in nitrogen content, but not water-soluble carbohydrate content of experimental swards.

We conclude that predictions of simple depletion models are unlikely to explain the movements of herbivores between patches that differ in digestibility and nutrient content as well as in the quantity of foods available and that multi-currency models are a more appropriate means of predicting foraging behaviour of herbivores exhibiting partial feeding preferences.