Habitat structure and association with ungulates drive habitat selection and grouping behaviour of lesser rhea (Rhea pennata subsp. pennata)

Social prey species respond to predation risk by modifying habitat selection and grouping behaviour. These responses may depend on both actual predation risk (predator probability of occurrence) and/or on perceived predation risk associated with habitat structure. Other factors like food availability and co-occurrence with other species may also affect habitat selection and group formation. We analyse habitat selection and grouping behaviour (group size and cohesion) of lesser rhea (Rhea pennata subsp. pennata), a ratite endemic of South America inhabiting steppe shrublands and grasslands, in relation to actual (puma probability of occurrence) and perceived (habitat structure: openness, visibility) predation risk, co-occurrence with other herbivore species and forage availability in the Chilean Patagonia. We used data from 9 sampling seasons in 5 years. Results show that habitat selection, group size and cohesion in lesser rhea were mainly driven by variables associated with perceived predation risk and by co-occurrence with other herbivores both during breeding and non–breeding season. As expected, lesser rhea preferred open habitats (vegas and grasslands) that allow a behaviour of ‘watch and run’ to avoid predation and formed larger groups in them. Moreover, lesser rhea positively selected year-round habitats where livestock occur, forming large groups during non–breeding season there. Group size and co-occurrence with other herbivores significantly decreased group cohesion, suggesting a reduction of perceived predation risk. Therefore, lesser rhea seems to take advantage of forming mixed interspecific groups to reduce predation risk. These results suggest that lesser rhea habitat selection and grouping behaviour are preferentially driven by factors related to perceived predation risk than by actual predator occurrence or food availability.     

Food quality and quantity are more important in explaining foraging of an intermediate‐sized mammalian herbivore than predation risk or competition

During times of high activity by predators and competitors, herbivores may be forced to forage in patches of low‐quality food. However, the relative importance in determining where and what herbivores forage still remains unclear, especially for small‐ and intermediate‐sized herbivores. Our objective was to test the relative importance of predator and competitor activity, and forage quality and quantity on the proportion of time spent in a vegetation type and the proportion of time spent foraging by the intermediate‐sized herbivore European hare (Lepus europaeus). We studied red fox (Vulpes vulpes) as a predator species and European rabbit (Oryctolagus cuniculus) as a competitor. We investigated the time spent at a location and foraging time of hare using GPS with accelerometers. Forage quality and quantity were analyzed based on hand‐plucked samples of a selection of the locally most important plant species in the diet of hare. Predator activity and competitor activity were investigated using a network of camera traps. Hares spent a higher proportion of time in vegetation types that contained a higher percentage of fibers (i.e., NDF). Besides, hares spent a higher proportion of time in vegetation types that contained relatively low food quantity and quality of forage (i.e., high percentage of fibers) during days that foxes (Vulpes vulpes) were more active. Also during days that rabbits (Oryctolagus cuniculus) were more active, hares spent a higher proportion of time foraging in vegetation types that contained a relatively low quality of forage. Although predation risk affected space use and foraging behavior, and competition affected foraging behavior, our study shows that food quality and quantity more strongly affected space use and foraging behavior than predation risk or competition. It seems that we need to reconsider the relative importance of the landscape of food in a world of fear and competition.     

Interference competition and group size effect in sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) at salt licks

Competition has long been considered as a confounding factor of group size effect but the understanding of interference competition is rudimentary for the difficulty in disentangling interference competition from scramble competition adequately. Here, we analysed remote-camera video records of wild sika deer (Cervus nippon) at salt licks in southern China from March 1, 2006 to November 30, 2008 to investigate how interference competition and predation risk interacted on vigilance behaviour. Scramble competition is negligible at salt licks; therefore, we could focus our interest in interference competition. We used linear mixed model to compare vigilance, licking and aggression behaviours among females with and without fawn as well as males with different group sizes to identify the primary role of vigilance behaviour in sika deer. In total, 168 individuals were recorded and observation time was 2,733.04 min. We found that deer spent much time on vigilance and scanned frequently in spring and winter, and females with fawn spent more time on vigilance than females without fawn, suggesting vigilance for predation risk. Aggression ratio increased first and then decreased, while scan frequency continued to decline and then slightly increased when group size increased from two to seven, implying vigilance for interference competition. Our results suggested vigilance in sika deer was influenced by both predation risk and interference competition, but was mainly driven by predation risk even at sites with intense interference competition. Our results of interference competition shed some light on finding the underlying mechanism of group size effect in wild populations.     

Do elephants prevent other African herbivores from using waterholes in the dry season?

In some African protected areas, concerns have arisen about the influence of locally high elephant numbers on other forms of biodiversity. In arid and semi-arid savannas, surface-water resources are scarce and agonistic interactions between elephants and other herbivores have been reported at waterholes, yet surprisingly very little is known about the impact of elephants on the use of waterholes by other herbivores. Here, we test whether when there are elephants at a waterhole, other herbivores (1) do not change their drinking behaviour; (2) spend shorter time around the water because they are disturbed by elephants’ presence and consequently have to leave the waterhole area probably without having met their water requirements, or (3) spend more time around the water probably owing to an increase in vigilance activities or because the presence of elephants may signal safety from predators. Results show that all species spend longer time around water when there are elephants at the waterhole, although the difference is not large. Consequently, this study strongly suggests that elephants do not prevent other herbivores from drinking (time at waterholes is not shortened when elephants are around). Further, if the additional time spent to drink is linked to an increased vigilance, the difference is not large, and hence unlikely to affect the population dynamics of other herbivores.     

Behavioral responses of a sex-role reversed pipefish to a gradient of perceived predation risk

Conspicuous behaviors such as courtship and mating often makeanimals susceptible to predation. When perceiving themselvesat an elevated level of risk, animals frequently reduce conspicuousbehaviors in tradeoff for a decrease in probability of beingpreyed upon. In the present study, we used two experiments toexamine the effect of perceived predation risk from cod (Gadusmorhud) on nonreproductive and reproductive behaviors in thesex-role reversed pipefish (Syngnathus typhle). In the firstexperiment, no differences due to predation risk were detectedin the nonreproductive behaviors of either males or females.In the second experiment, predation risk had significant effectson reproductive behaviors. Pipefish were allowed to court andcopulate at four different predation levels. We created predationlevels differing in perceived predation risk by controllingthe number of sensory modes through which pipefish could detectthe presence of a cod. As predation risk increased, pipefishcopulated and courted less frequently, swam alone (displayedand searched for conspecifics) less often, and waited longerbefore commencing courtship. These changes in behavior minimizedthe amount of time spent above the eelgrass and presumably reducedconspicuousness to visual predators. Pipefish also copulatedafter a smaller amount of courtship as predation risk increased,indicating that they may trade information concerning mate qualityfor a reduction in predation risk. No differences were foundin any response variable between males and females. The roleof operational sex ratios and intersexual competition in determiningwhich sex assumes greater costs in mate acquisition is questioned.     

Large herbivore assemblages in a changing climate: incorporating water dependence and thermoregulation

The coexistence of different species of large herbivores (ungulates) in grasslands and savannas has fascinated ecologists for decades. However, changes in climate, land‐use and trophic structure of ecosystems increasingly jeopardise the persistence of such diverse assemblages. Body size has been used successfully to explain ungulate niche differentiation with regard to food requirements and predation sensitivity. But this single trait axis insufficiently captures interspecific differences in water requirements and thermoregulatory capacity and thus sensitivity to climate change. Here, we develop a two‐dimensional trait space of body size and minimum dung moisture content that characterises the combined food and water requirements of large herbivores. From this, we predict that increased spatial homogeneity in water availability in drylands reduces the number of ungulate species that will coexist. But we also predict that extreme droughts will cause the larger, water‐dependent grazers as wildebeest, zebra and buffalo–dominant species in savanna ecosystems – to be replaced by smaller, less water‐dependent species. Subsequently, we explore how other constraints such as predation risk and thermoregulation are connected to this two‐dimensional framework. Our novel framework integrates multiple simultaneous stressors for herbivores and yields an extensive set of testable hypotheses about the expected changes in large herbivore community composition following climate change.     

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.     

Dietary specialization is conditionally associated with increased ant predation risk in a temperate forest caterpillar community

The enemy‐free space hypothesis (EFSH) contends that generalist predators select for dietary specialization in insect herbivores. At a community level, the EFSH predicts that dietary specialization reduces predation risk, and this pattern has been found in several studies addressing the impact of individual predator taxa or guilds. However, predation at a community level is also subject to combinatorial effects of multiple‐predator types, raising the question of how so‐called multiple‐predator effects relate to dietary specialization in insect herbivores. Here, we test the EFSH with a field experiment quantifying ant predation risk to insect herbivores (caterpillars) with and without the combined predation effects of birds. Assessing a community of 20 caterpillar species, we use model selection in a phylogenetic comparative framework to identify the caterpillar traits that best predict the risk of ant predation. A caterpillar species' abundance, dietary specialization, and behavioral defenses were important predictors of its ant predation risk. Abundant caterpillar species had increased risk of ant predation irrespective of bird predation. Caterpillar species with broad diet breadth and behavioral responsiveness to attack had reduced ant predation risk, but these ant effects only occurred when birds also had access to the caterpillar community. These findings suggest that ant predation of caterpillar species is density‐ or frequency‐dependent, that ants and birds may impose countervailing selection on dietary specialization within the same herbivore community, and that contingent effects of multiple predators may generate behaviorally mediated life‐history trade‐offs associated with herbivore diet breadth.     

Costs of fear: behavioural and life‐history responses to risk and their demographic consequences vary across species

Behavioural responses to reduce predation risk might cause demographic ‘costs of fear’. Costs differ among species, but a conceptual framework to understand this variation is lacking. We use a life‐history framework to tie together diverse traits and life stages to better understand interspecific variation in responses and costs. We used natural and experimental variation in predation risk to test phenotypic responses and associated demographic costs for 10 songbird species. Responses such as increased parental attentiveness yielded reduced development time and created benefits such as reduced predation probability. Yet, responses to increased risk also created demographic costs by reducing offspring production in the absence of direct predation. This cost of fear varied widely across species, but predictably with the probability of repeat breeding. Use of a life‐history framework can aid our understanding of potential demographic costs from predation, both from responses to perceived risk and from direct predation mortality.     

Influence of predation risk on individual competitive ability and growth in Eurasian perch, Perca fluviatilis

We studied the potential influence of predation risk on the competitive ability and habitat use of foraging perch and the effect of these interactions on growth. Groups of four similar-sized young-of-the-year perch were in visual contact with a piscivorous perch during feeding. The fry had the choice of vegetation and open habitat, with food presented in the open habitat. Competitive ability, defined as proportion of prey attacks, varied between perch individuals and was unaffected by predation risk. The variation in proportion of prey attacks was affected by relative size within each replicate group, despite small size differences (±1 mm), with the largest individual being a better competitor than the smallest ones. The degree of boldness, measured as the proportion of time spent in the open habitat, was significantly related to both competitive ability and prey attack order. Observations of aggressive behaviour indicated a possible occurrence of interference competition, which may contribute to the appearance of different competitive abilities between individuals within a group of perch. A significant correlation was found between competitive ability and growth. Growth variation within groups was not affected by predation risk.     

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.     

Titrating the cost of plant toxins against predators: determining the tipping point for foraging herbivores

1. Foraging herbivores must deal with plant characteristics that inhibit feeding and they must avoid being eaten. Principally, toxins limit food intake, while predation risk alters how long animals are prepared to harvest resources. Each of these factors strongly affects how herbivores use food patches, and both constraints can pose immediate proximate costs and long-term consequences to fitness. 2. Using a generalist mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), our aim was to quantitatively compare the influence of plant toxin and predation risk on foraging decisions. 3. We performed a titration experiment by offering animals a choice between non-toxic food at a risky patch paired with food with one of five toxin concentrations at a safe patch. This allowed us to identify the tipping point, where the cost of toxin in the safe food patch was equivalent to the perceived predation risk in the alternative patch. 4. At low toxin concentration, animals ate more from the safe than the risky patch. As toxin concentration increased at the safe patch, intake shifted until animals ate mainly from the risky patch. This shift was associated with behavioural changes: animals spent more time and fed longer at the risky patch, while vigilance increased at both risky and safe patches. 5. Our results demonstrate that the variation in toxin concentration, which occurs intraspecifically among plants, can critically influence the relative cost of predation risk on foraging. We show that herbivores quantify, compare and balance these two different but proximate costs, altering their foraging patterns in the process. This has potential ecological and evolutionary implications for the production of plant defence compounds in relation to spatial variation in predation risk to herbivores.     

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.     

Behaviourally mediated indirect effects: interference competition increases predation mortality in foraging redshanks

1. The effect of competition for a limiting resource on the population dynamics of competitors is usually assumed to operate directly through starvation, yet may also affect survival indirectly through behaviourally mediated effects that affect risk of predation. Thus, competition can affect more than two trophic levels, and we aim here to provide an example of this. 2. We show that the foraging success of redshanks Tringa totanus (L.) foraging on active prey was highest in the front of flocks, whereas this was not the case for redshanks foraging on inactive prey. Also, when foraging on active prey, foraging success in a flock decreased as more birds passed through a patch, while overall foraging success was not lower on subsequent visits to the same patch. Thus, redshanks foraging on active prey suffered from interference competition, whereas this was not the case for redshanks foraging on inactive prey. 3. This interference competition led to differences in activity: redshanks attaining a lower foraging success had a higher walking rate. Greater activity was associated with wider flock spacing and shorter distances to cover, which has previously been shown to increase predation risk and mortality from sparrowhawks Accipiter nisus (L.). 4. We conclude that behavioural adaptations of prey species can lead to interference competition in foraging redshanks, and thus can affect their predation risk and mortality through increased activity. This study is one of the first to show how interference competition can be a mechanism for behaviourally mediated indirect effects, and provides further evidence for the suggestion that a single species occupying an intermediate trophic level may be simultaneously top-down controlled by a predator and bottom-up controlled by a behavioural response of its prey.     

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.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.     

Limited spatial response to direct predation risk by African herbivores following predator reintroduction

Predators affect ecosystems not only through direct mortality of prey, but also through risk effects on prey behavior, which can exert strong influences on ecosystem function and prey fitness. However, how functionally different prey species respond to predation risk and how prey strategies vary across ecosystems and in response to predator reintroduction are poorly understood. We investigated the spatial distributions of six African herbivores varying in foraging strategy and body size in response to environmental factors and direct predation risk by recently reintroduced lions in the thicket biome of the Addo Elephant National Park, South Africa, using camera trap surveys, GPS telemetry, kill site locations and Light Detection and Ranging. Spatial distributions of all species, apart from buffalo, were driven primarily by environmental factors, with limited responses to direct predation risk. Responses to predation risk were instead indirect, with species distributions driven by environmental factors, and diel patterns being particularly pronounced. Grazers were more responsive to the measured variables than browsers, with more observations in open areas. Terrain ruggedness was a stronger predictor of browser distributions than was vegetation density. Buffalo was the only species to respond to predator encounter risk, avoiding areas with higher lion utilization. Buffalo therefore behaved in similar ways to when lions were absent from the study area. Our results suggest that direct predation risk effects are relatively weak when predator densities are low and the time since reintroduction is short and emphasize the need for robust, long‐term monitoring of predator reintroductions to place such events in the broader context of predation risk effects.     

Predation risk for herbivorous insects on tropical vegetation: A search for enemy-free space and time

Spatial and temporal variability in predation risk for herbivores on 13 rainforest species of Ficus (Moraceae) in Papua New Guinea was studied in order to assess whether predator-free refuges exist on their foliage and if so, whether herbivorous insects concentrate their activity in such refugia. Predation risk from invertebrate predators was measured as the disappearance rate of live termites set up as baits on the foliage. By far the most important predators were ants, accounting for 77% of attacks. No consistent differences in predation rate between Ficus species were found so that tree identity could not be used as an indicator of enemy-free space. Predation risk was highly variable among conspecific trees and also changed rapidly in time, over periods as short as 10 days. Such short-term and unpredictable predator-free refuges may be difficult for herbivores to find and exploit. Predation risk during the day was three times higher than during the night, but abundance of herbivores on the foliage was also higher during the day. Thus, night was confirmed as a relatively enemy-free time which, however, was not exploited by herbivores.     

Interference competition and temporal niche shifts: elephants and herbivore communities at waterholes

Scarcity of resources may result in high levels of animal aggregation; interference competition can occur in such a scenario and play a role in resource acquisition. Here, we test the hypothesis that animals could minimize interference competition by shifting their temporal niches in relation to competitors. In Hwange National Park, Zimbabwe, we monitored waterholes in order to study agonistic interactions between elephants and other herbivore species. We also used a long-term data set from a yearly survey of waterhole attendance by herbivores to evaluate the influence of the presence of elephants on the use of waterholes by other herbivore species. Our results show that in drier years, waterholes are crowded with elephants early in the afternoon. In general, the species most affected by interference competition with elephants shift their temporal niches at the waterholes, thus maintaining a constant temporal overlaps with elephants. The species less affected by interference competition with elephants show no temporal niche shifts and increase their temporal overlap with elephants at waterholes, as predicted from a noncompetition hypothesis. This study provides evidence that interference competition with a behaviorally dominant large species influences the temporal niches of smaller species, and suggests that the potential costs associated with interference between elephants and other herbivores at waterholes are linked to shifts in diurnal activities rather than interactions and water acquisition itself.     

Nutrient-limited food webs with up to three trophic levels: feasibility, stability, assembly rules, and effects of nutrient enrichment

Community structure is controlled, among multiple factors, by competition and predation. Using the R* rule and graphical analysis, we analyse here the feasibility, stability and assembly rules of resource-based food webs with up to three trophic levels. In particular, we show that (1) the stability of a food web with two plants and two generalist herbivores does not require that plants' resource exploitation abilities trade-off with resistance to the two herbivores, and (2) food webs with two plants and either one generalist herbivore and a carnivore or two generalist herbivores and two generalist carnivores are not feasible because of cascade competition between top consumers. The relative strength of species interactions and the relative impacts of plants and herbivores on factors which control their growth also play a critical role. We discuss how community structure constrains assembly rules and yields cascades of extinctions in food webs.