Olfactory misinformation provides refuge to palatable plants from mammalian browsing.

Mammalian herbivores browse palatable plants of ecological and economical value. Undesirable neighbours can reduce browsing to these plants by providing 'associational refuge', but they can also compete for resources. Here we recreated the informative odour emitted by undesirable plants. We then tested whether this odour could act as virtual neighbours, providing browsing refuge to palatable eucalyptus tree seedlings. We found that protection using this method was equivalent to protection provided by real plants. Palatable seedlings were 17-20 times more likely to be eaten by herbivores without virtual, or real, neighbours. Because many herbivores use plant odour to forage, virtual neighbours could provide a useful practical management approach to help protect valued plants.

Ecology: Humans are scarier than lions.

In Africa, nothing inspires fear more than lions. They are large, hunt in groups and kill prey much larger than themselves. Yet, evidence suggests that African wildlife are more afraid of humans than anything else that moves across the savanna.

The deterrent effects of individual monoterpene odours on the dietary decisions of African elephants.

African savanna elephants use pre-ingestive olfactory cues when making dietary choices, and previous research has observed that elephant diet choice is negatively correlated with vegetation species that contain high concentrations of monoterpenes. However, the frequency and concentration of monoterpenes can vary dramatically across plant species. Thus, we aimed to explore the effects that the odours of individual monoterpenes have on elephant diet choice and how these effects vary with concentration. To do this, we conducted three odour-based choice experiments focusing on eight common monoterpenes found in the woody plants in Southern African savannas. In the first experiment, we tested whether elephant diet choice for a frequently consumed plant (Euclea crispa) was influenced by the addition of the odour of an individual monoterpene at a set concentration. In the second experiment, we explored the relative deterrence of each monoterpene. Lastly, we tested how elephant diet choice varied as a function of the addition of individual monoterpene odours at 5%, 10%, and 20% concentrations. We found that the elephants avoided most individual monoterpenes at high concentrations, with the exception being α-pinene. Furthermore, we found that the odours of some individual monoterpenes were, in fact, more deterrent than others. In the third experiment, we found that the elephants avoided ß-pinene, limonene, ocimene, γ-terpinene, and terpinolene across all concentrations, but only avoided sabinene and linalool at high concentrations. Ultimately, our results show that the odour of individual monoterpenes may deter elephant consumption, but that this deterrent effect depends on both the monoterpene and its concentration.

The Olfactory Landscape Concept: A Key Source of Past, Present, and Future Information Driving Animal Movement and Decision-making.

Odor is everywhere, emitted across the landscape from predators, prey, decaying carcasses, conspecifics, vegetation, surface water, and smoke. Many animals exploit odor to find food, avoid threats, and attract or judge potential mates. Here, we focus on odor in terrestrial ecosystems to introduce the concept of an olfactory landscape: real-time dynamic olfactory contours reflecting the patchy distribution of resources and risks, providing a key source of information used by many animals in their movement and decision-making. Incorporating the olfactory landscape into current frameworks of movement ecology and animal behavior will provide a mechanistic link to help answer significant questions about where, why, and when many animals move, and how they do so efficiently in both space and time. By understanding how animals use the olfactory landscape to make crucial decisions affecting their fitness, we can then manipulate the landscape to modify ecological interactions and, ultimately, ecosystem consequences of these interactions.

Cueing on distant conditions before migrating does not prevent false starts: a case study with African elephants.

Migratory animals often use environmental cues to time their seasonal migrations. Local conditions may, however, differ from distant ones, and current conditions may poorly predict future conditions. This may be particularly true for early wet season conditions in tropical systems, as storms and associated rainfall events are generally not predictable at the scale of weeks or days and are heterogeneously distributed even at the scale of a few kilometres. How migratory animals cope with such challenges, and the consequences they may have, remain poorly known. We used time-to-event models based on GPS data from 19 African elephant herds from Hwange National Park (Zimbabwe) to study the effect of local and distant rainfall events on the elephants' decision to initiate their wet season migration. Elephants relied more on distant rainfall events occurring along the future migration route than on local events when initiating their migration. Such ability to use distant cues does not, however, ensure an immediate migration success. In over 30% of the cases, the elephants came back to their dry season range, sometimes after having travelled > 80% of the expected migration distance. This happened particularly when there was little additional rain falling during the migration. All elephants successfully migrated later in the season. Our study improves the understanding of the migratory ecology of elephants. More broadly, it raises questions about the reliability of rainfall as a migratory cue in tropical systems, and shed light on one of its potential consequences, the poorly quantified phenomenon of migration false starts.

Individuality, species-specific features, and female discrimination of male southern white rhinoceros courtship calls.

Male vocalizations associated with courtship can play a key role in mate selection. They may help females obtain information about males' quality and identity and/or may contain species-specific properties that help prevent interspecies breeding. Despite vocalizations being a prominent part of the courtship of white rhinos, the role that they play in white rhino breeding behaviour has not been extensively studied. Both southern (SWR) and critically endangered northern white rhino (NWR) males intensively vocalize during courtship with hic calls. We examined these calls and found that call properties differed between NWR and SWR males. In addition, we found that individual SWR males could be identified with a high degree of accuracy using their hic calls and that the signature information capacity in hic calls would allow females to individually recognize about 11 adult males living in or moving through their home-ranges, which may help with mate selection. Then, we conducted playback experiments with wild anoestrus SWR females. The females discriminated between the NWR and SWR hic calls and between the SWR hic and SWR pant calls. However, we only found differences in the latency of observed behaviours, not in their duration or in the intensity of females' reaction. This might suggest that females which are not in oestrus are not highly responsive to a male's motivation (i.e., seeking contact or mating), but are more interested in assessing his dominance status or familiarity. Ultimately, our results indicate that courtship hic calls encode information which might help females choose mating partners.

African elephants can detect water from natural and artificial sources via olfactory cues.

Water is vital for mammals. Yet, as ephemeral sources can be difficult to find, it raises the question, how do mammals locate water? Elephants (Loxodonta africana) are water-dependent herbivores that possess exceptional olfactory capabilities, and it has been suggested that they may locate water via smell. However, there is no evidence to support this claim. To explore this, we performed two olfactory choice experiments with semi-tame elephants. In the first, we tested whether elephants could locate water using olfactory cues alone. For this, we used water from two natural dams and a drinking trough utilised by the elephants. Distilled water acted as a control. In the second, we explored whether elephants could detect three key volatile organic compounds (VOCs) commonly associated with water (geosmin, 2-methylisoborneol, and dimethyl sulphide). We found that the elephants could locate water olfactorily, but not the distilled water. Moreover, they were also able to detect the three VOCs associated with water. However, these VOCs were not in the odour profiles of the water sources in our experiments. This suggests that the elephants were either able to detect the unique odour profiles of the different water sources or used other VOCs that they associate with water. Ultimately, our findings indicate that elephants can locate water olfactorily at small spatial scales, but the extent to which they, and other mammals, can detect water over larger scales (e.g. km) remains unclear.

Are there phylogenetic differences in salivary tannin-binding proteins between browsers and grazers, and ruminants and hindgut fermenters?

While feeding, mammalian browsers (primarily eat woody plants) encounter secondary metabolites such as tannins. Browsers may bind these tannins using salivary proteins, whereas mammalian grazers (primarily eat grasses that generally lack tannins) likely would not. Ruminant browsers rechew their food (ruminate) to increase the effectiveness of digestion, which may make them more effective at binding tannins than nonruminants. Few studies have included a sufficient number of species to consider possible scaling with body mass or phylogenetic effects on salivary proteins. Controlling for phylogeny, we ran inhibition radial diffusion assays of the saliva of 28 species of African herbivores that varied in size, feeding strategy, and digestive system. We could not detect the presence of salivary proline-rich proteins that bind tannins in any of these species. However, using the inhibition radial diffusion assay, we found considerable abilities to cope with tannins in all species, albeit to varying degrees. We found no differences between browsers and grazers in the effectiveness of their salivary proteins to bind to and precipitate tannins, nor between ruminants and nonruminants, or scaling with body mass. Three species bound all tannins, but their feeding niches included one browser (gray duiker), one mixed feeder (bush pig), and one grazer (red hartebeest). Five closely related species of small ruminant browsers were very effective in binding tannins. Megaherbivores, considered generalists on account of their large body size, were capable of binding tannins. However, the grazing white rhinoceros was almost as effective at binding tannins as the megaherbivore browsers. We conclude, contrary to earlier predictions, that there were no differences in the relative salivary tannin-binding capability that was related to common ancestry (phylogeny) or to differences in body size.

Megaherbivore browsers vs. tannins: is being big enough?

Megaherbivores have been of particular interest to scientists because of the physiological and ecological challenges associated with their extreme body size. Yet, one question that has seldom been explored is how browsing megaherbivores cope with plant secondary metabolites (PSMs), such as tannins, found in their food. It is possible that the sheer body size of these megaherbivores allows them to ingest tannins with no deleterious effects. However, it is plausible that megaherbivores must rely on other mechanisms to cope with PSMs, such as the production of salivary tannin-binding proteins. Thus, we aimed to determine whether megaherbivore browsers produce tannin-binding proteins to further reduce the consequences of ingesting a tannin-rich diet. Using a series of laboratory assays, we explored whether elephants, black rhinoceros, and giraffe had tannin-binding proteins in their saliva. We tested for the presence of proline-rich proteins in the saliva using two different approaches: (1) SDS-PAGE using Laemmli's (Laemmli, Nature 227:680-685, 1970) destaining method, and (2) comparative SDS-PAGE gels using Beeley et al.'s (Beeley et al. Electrophoresis 12:493-499, 1991) method for staining and destaining to probe for proline-rich proteins. Then, to test for the tannin-binding affinity of their saliva, we performed an inhibition assay. We did not observe proline-rich proteins in any of the megaherbivore species, but they did have other protein(s) in their saliva that have a high tannin-binding affinity. Our results highlight that, despite their large body sizes, and their abilities to tolerate low-quality food, browsing megaherbivores have likely evolved tannin-binding proteins as a way of coping with the negative effects of tannins.

Coexistence between wildlife and livestock is contingent on cattle density and season but not differences in body size.

Many studies on the coexistence of wildlife with livestock have focused primarily on similar-sized species. Furthermore, many of these studies have used dietary overlap as a measure of potential competition between interacting species and thus lack the important link between dietary overlap and any negative effects on a particular species-a prerequisite for competition. Consequently, the mechanisms that drive interspecific interactions between wildlife and cattle are frequently overlooked. To address this, we used an experimental setup where we leveraged different cattle stocking rates across two seasons to identify the drivers of interspecific interactions (i.e. competition and facilitation) between smaller-bodied oribi antelope and cattle. Using direct foraging observations, we assessed dietary overlap and grass regrowth, and also calculated oribi nutritional intake rates. Ultimately, we found that cattle compete with, and facilitate, smaller-bodied oribi antelope through bottom-up control. Specifically, cattle facilitated oribi during the wet season, irrespective of cattle stocking density, because cattle foraging produced high-quality grass regrowth. In contrast, during the dry season, cattle and oribi did not co-exist in the same areas (i.e. no direct dietary overlap). Despite this, we found that cattle foraging at high densities during the previous wet season reduced the dry season availability of oribi's preferred grass species. To compensate, oribi expanded their dry season diet breadth and included less palatable grass species, ultimately reducing their nutritional intake rates. Thus, cattle competed with oribi through a delayed, across-season habitat modification. We show that differences in body size alone may not be able to offset competitive interactions between cattle and wildlife. Finally, understanding the mechanisms that drive facilitation and competition are key to promoting co-existence between cattle and wildlife.

Mixed-species herding levels the landscape of fear.

Prey anti-predator behaviours are influenced by perceived predation risk in a landscape and social information gleaned from herd mates regarding predation risk. It is well documented that high-quality social information about risk can come from heterospecific herd mates. Here, we integrate social information with the landscape of fear to quantify how these landscapes are modified by mixed-species herding. To do this, we investigated zebra vigilance in single- and mixed-species herds across different levels of predation risk (lion versus no lion), and assessed how they manage herd size and the competition-information trade-off associated with grouping behaviour. Overall, zebra performed higher vigilance in high-risk areas. However, mixed-species herding reduced vigilance levels. We estimate that zebra in single-species herds would have to feed for approximately 35 min more per day in low-risk areas and approximately 51 min more in high-risk areas to compensate for the cost of higher vigilance. Furthermore, zebra benefitted from the competition-information trade-off by increasing the number of heterospecifics while keeping the number of zebra in a herd constant. Ultimately, we show that mixed-species herding reduces the effects of predation risk, whereby zebra in mixed-species herds, under high predation risk, perform similar levels of vigilance compared with zebra in low-risk scenarios.

Shrub cover homogenizes small mammals' activity and perceived predation risk.

Altered disturbance regimes, increasing atmospheric CO2, and other processes have increased woody cover and homogenized vegetation in savannas across the planet. African savannas with extensive versus minimal woody cover often have vastly different animal communities. However, we lack a clear mechanistic understanding of why animal communities are changing with vegetation structure. Our goal for this study was to understand how vegetation structure in an African savanna shaped the perceived predation risk of small mammals, hence affecting their activity. Using a reciprocal measure of standard giving-up-densities, amount of food eaten, we found sharp declines in rodents' perceived predation risk and increased rodent activity underneath shrub cover. This response was consistent across species; however, species showed subtle differences in their responses to grassy vegetation. Our findings suggest that areas of minimal or extensive shrub cover (shrub encroachment) may be homogenizing rodents' perceptions of predation risk and thus shaping their use of space.

Author Correction: Temporal Variation of White Rhino Dung Odours.

The original version of this article unfortunately contained some mistakes.

Alarm calls or predator calls: which elicit stronger responses in ungulate communities living with and without lions?

Alarm calls and predator vocalizations convey information on predator presence and potential risk. Generally, prey employ anti-predator behaviours more in response to alarm calls. However, occasionally prey respond more to the vocalizations of specific predators. A key question is do prey still respond to alarm calls and predator vocalizations when a dangerous predator is absent? Additionally, would the prey species' response (e.g. vigilance) differ from prey already living with these predators? Using auditory playbacks, we tested whether four herbivore species living with lions responded more to alarm calls than lion vocalizations compared to a black cuckoo control call. Overall, red hartebeest, wildebeest and zebra had greater vigilance in response to the lion roars compared to the alarm calls. The differences in vigilance suggest that, despite the lion roars not being related to hunting, these herbivores perceived the predator vocalizations as a more immediate indicator of risk than the alarm calls. We then tested whether herbivores living with lions increased their vigilance more in response to the calls than conspecifics in a lion-free section. Despite greater overall vigilance in the lion section, gemsbok and zebra in the lion-free section significantly increased their vigilance in response to the lion roars. This indicates that species under the greatest threat from a predator (e.g. preferred prey) may maintain innate anti-predator responses to an absent but dangerous predator longer than less preferred prey. Ultimately, our results indicate that cues from dangerous predators can have greater effects on anti-predator behaviours than alarm calls for some prey species.

Can intrinsic foraging efficiency explain dominance status? A test with functional response experiments.

The functional response describes how food abundance affects the intake rate of foraging individuals, and as such, it can influence a wide range of ecological processes. In social species, dominance status can affect the functional response of competing individuals, but studies conducted in an interference-free context have provided contrasting results on the extent of between-individual variability in functional response. We tested the prediction that individuals intrinsically differ in their functional response, and that these differences could predict body weight and dominance status in social species. We used goats as a model species and performed foraging experiments to assess the functional response of these goats in an interference-free context. Our results show that some individuals are consistently better foragers than others, and these individuals were more likely to be heavier and dominant. Parameters of the functional response are, however, more strongly associated with dominance status than with body weight. We conclude that interference while foraging is not needed to explain body weight differences between dominant and subordinate individuals. We suggest that these differences can emerge from intrinsic differences in foraging efficiency between individuals, which could also allow better foragers to demonstrate greater tenacity during agonistic interactions.

Temporal Variation of White Rhino Dung Odours.

In order for an olfactory signal to be effective, it must persist in the environment for an extended period. White rhino dung odours transmit information about sex, age, territorial and oestrous states. As these odours relay important information, temporal changes in the odour emission rate and/or composition may be critical in order for other individuals to obtain this information. Here, we examine how the dung odours of adult white rhinos (male: territorial and non-territorial; female: oestrous and non-oestrous) change over the short (hours) and long (seasons) term using headspace extraction. Additionally, we measure seasonal midden visitation and defecation frequency to link behaviours to seasonal changes in odour longevity. We found that during the dry season, territorial male dung odours persisted for 48 hours, while the odours of all other adults persisted for >48 hours. The high temperature and humidity of the wet season did not affect odour longevity of oestrous females, but decreased dung odour longevity (i.e. increased emission) of territorial males, non-territorial males, and non-oestrous females. Despite this reduction, neither males nor females adjusted their seasonal visitation or defecation frequency. With regard to a temporal indicator, 3,7-dimethly-1,6-octadiene signalled dung age during the wet season, while acetophenone signalled dung age during the dry season. Ultimately, our results provide the first detailed account of temporal changes of dung VOCs, and highlight the limitations of dung as a signalling medium.

Dung odours signal sex, age, territorial and oestrous state in white rhinos.

Mammals commonly communicate olfactorily via urine. However, the extent to which they communicate via dung, another waste product, is unknown. Behavioural studies suggest that mammals can obtain information from dung odours but are unclear about the information transmitted. Moreover, an understanding of the volatile organic compounds (VOCs) released from dung is limited. To address this, we analysed the odours emitted from the dung of free-ranging white rhinos, and found that 2,3-dimethylundecane signalled an individual's sex, heptanal discriminated age class, nonane defined male territorial status and 2,6-dimethylundecane indicated female oestrous state. To validate these findings, we artificially reproduced key elements of the territorial and oestrous odour profiles (i.e. profiles likely to elicit behavioural responses from receivers). We then exposed free-ranging territorial males to these odours. In response, males elicited behaviours associated with the specific odours (e.g. territorial male (potential threat): reduced latency in assuming vigilance; oestrous female (potential mate): increased investigation). These results indicate that the VOCs identified from the dung of free-ranging individuals do transmit key information. Moreover, as white rhinos of all ages and sexes defecate communally, middens probably act as information centres. Furthermore, as many other mammals defecate communally, olfactory communication via dung odours is likely a widespread phenomenon.

Group-living herbivores weigh up food availability and dominance status when making patch-joining decisions.

Two key factors that influence the foraging behaviour of group-living herbivores are food availability and individual dominance status. Yet, how the combination of these factors influences the patch-joining decisions of individuals foraging within groups has scarcely been explored. To address this, we focused on the patch-joining decisions of group-living domestic goats (Capra hircus). When individuals were tested against the top four ranked goats of the herd, we found that at patches with low food availability they avoided these dominant patch-holders and only joined subordinates (i.e. costs outweighed benefits). However, as the amount of food increased, the avoidance of the top ranked individuals declined. Specifically, goats shifted and joined the patch of an individual one dominance rank higher than the previous dominant patch holder when the initial quantity of food in the new patch was twice that of the lower ranking individual's patch (i.e. benefits outweighed costs). In contrast, when individuals chose between patches held by dominant goats, other than the top four ranked goats, and subordinate individuals, we found that they equally joined the dominant and subordinate patch-holders. This joining was irrespective of the dominance gap, absolute rank of the dominant patch-holder, sex or food availability (i.e. benefits outweighed costs). Ultimately, our results highlight that herbivores weigh up the costs and benefits of both food availability and patch-holder dominance status when making patch-joining decisions. Furthermore, as the initial quantity of food increases, food availability becomes more important than dominance with regard to influencing patch-joining decisions.