Background: Browsing repellents are widely used to deter large herbivores from consuming plants of ecological,economic and aesthetic importance. Understanding how these repellents function on a behavioural mechanisticlevel is critical to predicting effectiveness. Here, we illustrate how these mechanisms can be tested, by exposing amodel mammalian herbivore, the fallow deer, to different concentrations of a commercial chemical repellent(HaTe2) in two-choice feeding trials.

Results:The repellent acted as a defensive chemical for the food by both reducing visitation and the amountconsumed. Deer favoured the less defended feeders before ingesting any food, suggesting that the repellentaltered olfactory and/or visual cues. Deer also consumed less of the more defended food when choosing betweenlow and high repellent feeders than no and low repellent feeders, indicating that the repellent modified flavourand/or sensation. Repellent effectiveness declined with increased exposure, suggesting that consumption had nonegative post-ingestive effects, and thus, deterrence was not caused by a conditioned aversion or irritation. Instead,this pattern suggests that deer learned, through repeated sampling of repellent-treated food, that there was noadverse physiological effect of ingesting it.

Conclusions:These results imply that HaTe2 repellent will not be effective over prolonged periods or in theabsence of alternative untreated food. Understanding the mechanisms driving repellent function using two-choicetrials could help practitioners decide whether a particular repellent is likely to be effective against mammalianherbivory in their management scenario.

Browsing can reduce forest productivity, particularly when the apical shoots of trees are damaged. Repellents are used widely to reduce browsing, but application is costly. To improve efficiency, it may be possible to take advantage of associational plant refuge effects, requiring repellents to be applied only to some trees or parts of trees, or reapplied less frequently. Using captive moose (Alces alces) and constructed stands of Scots pine (Pinus sylvestris), we tested for potential refuges by applying a commercial repellent (HaTe2) to all, alternate or none of the apical shoots, or all of the previous-year apical shoots. We also tested for potential refuges under field conditions, applying the repellent to all, alternate or none of the apical pine shoots in forest stands. Captive moose (two individuals in a 2.1-ha enclosure, similar to 95 individuals km(-2)) browsed 100% of trees, but were significantly less likely to browse apical shoots treated with repellent. Associational refuge was ineffective both within and between trees. In the field (0.84 moose km(-2)), only 1.3% of trees sustained browsing damage. Applying the repellent to the apical shoots of pines had no direct repellent effect nor any within-plant associational effects. Trees with treated apical shoots provided some protection for untreated neighbouring trees, but this was not biologically meaningful given the low percentage of trees browsed overall. Here, a simple captive experiment was not predictive of the browsing response observed in the field, demonstrating the need to test repellent application strategies in situ.

For social foragers, movement as a group could increase foraging efficiency through collective discovery of high-quality food sources. This would require an efficient mechanism for transferring information about food quality between individuals. Conversely, the constraints of foraging as a cohesive group could decrease efficiency; grouping may persist to serve other functions such as protection from predators. To test what drives cohesion in herbivores, we manipulated patch shape and within-patch pattern of food quality and quantified the effects on group level diet selection by a social herbivore, the fallow deer, Dama dama. We arranged feeders containing fodder in lines or blocks, and manipulated the pattern of food quality within patches by adding tannin, a plant secondary compound that decreases palatability. We quantified the relative consumption of low- and high-tannin food to compare diet selectivity at the group level between patch treatments. If group foraging evolved to increase foraging efficiency, altering the spatial arrangement of food should not affect diet selectivity because information about food location and quality is shared. We found, however, that the herd expressed different levels of selectivity between both patch shapes and food quality patterns. Deer selected better diets in blocks than lines. In lines, the herd selected better diets when quality varied between alternate feeders rather than between the two halves of the patch, suggesting a reliance on personal rather than group information. Deer consumed the most at patch centres in all treatments except in blocks with high-tannin centres, but diet selection was poorer in the latter compared to blocks with low-tannin centres. Aggregation at the centre of patches appears to have restricted exploitation of the best food. Predation pressure and/or resource variability may have favoured the evolution of a foraging strategy that prioritizes social cohesion over effective diet selection.

Mammalian browsers can cause ecologically and economically significant damage to important tree species, particularly at the vulnerable seedling stage. Some tree species or particular parts of trees are less preferred as a result of chemical defences - harnessing such defences could reduce the appeal of more preferred trees and thus their loss to browsing. We tested the potential of birch bark extract, a forestry by-product, to protect seedlings of Scots pine Pinus sylvestris from two globally significant browsers: moose Alces alces and red deer Cervus elaphus. We quantified the responses by captive animals to extract application as a function of both the distance between seedlings and whether the extract was applied to all or alternate seedlings. Both moose and red deer consumed pine seedlings less frequently when seedlings were treated with birch bark extract, but their responses differed when only alternate seedlings were treated. At inter-seedling distances equivalent to those used in forestry, red deer browsed untreated seedlings more frequently when alternate seedlings were treated than when none were treated (associational susceptibility), whereas moose browsed untreated seedlings less frequently with treated than untreated neighbours (associational refuge). These neighbourhood effects were not evident at inter-seedling distances three times larger or smaller than those used in forestry. There was also no significant difference in the frequency of browsing on treated seedlings in alternate compared to completely treated arrangements at any inter-seedling distance. Red deer removed significantly less foliage from treated than untreated seedlings once browsed. Browsing by moose followed a similar but non-significant trend. The relative number of untreated to treated seedlings browsed was 2:1 for red deer and 4:1 for moose. Red deer used a finer scale of selection than moose for choosing between seedlings, consistent with their smaller body mass. However, the coarser-scale of selection used by moose resulted in consumption of more untreated relative to treated seedlings. Our study highlights the potential of existing plant defensive chemistry, harvested from forestry by-products, to reduce herbivore damage to tree seedlings. We also demonstrate the need to consider both plant associational effects and the foraging strategies of the dominant herbivores in applying repellents.

Searching for food is the first critical stage of foraging, and search efficiency is enhanced when foragers use cues from foods they seek. Yet we know little about food cues used by one major group of mammals, the herbivores, a highly interactive component of most ecosystems. How herbivores forage and what disrupts this process, both have significant ecological and evolutionary consequences beyond the animals themselves. Our aim was to investigate how free-ranging mammalian herbivores exploit leaf odour cues to find food plants amongst a natural and complex vegetation community. Our study system comprised the native deer equivalent of eastern Australian forests, the swamp wallaby Wallabia bicolor, and seedlings of Eucalyptus, the foundation tree genus in these ecosystems. We quantified how foraging wallabies responded to odour cues from plants manipulated in several ways: varying the quantity of visually concealed leaves, comparing damaged vs. undamaged leaves, and whole plants vs. those with suppressed cues. The rate of discovery of leaves by wallabies increased with odour cue magnitude, yet animals were extremely sensitive to even a tiny odour source of just a few leaves. Whole seedlings were discovered faster if their leaves were damaged. Wallabies found whole seedlings and those with suppressed visual cues equally rapidly, day and night. Seedlings with very little odour were discovered mainly by day, as nocturnal foraging success was severely disrupted. This study shows how leaf odour attracts mammalian herbivores to food plants, enabling non-random search for even tiny odour sources. As damaged leaves enhanced discovery, we suggest that the benefit of attracting natural enemies to invertebrate herbivores feeding on plants (potential cry for help) may be offset by a cost-increased browsing by mammalian herbivores. This cost should be incorporated into multi-trophic plant-animal studies. Finally, the breakdown in capacity to find plants at night suggests substantial but unrecognized foraging costs to herbivores when abiotic factors, such as cold temperatures or pollution, reduce or degrade plant odour cues. We predict that an increasingly polluted world will alter the foraging success of mammalian herbivores, with significant ecological ramifications given that browsing can shape ecosystems.

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.

Studies of odour-driven foraging by mammals focus on attractant cues emitted by flowers, fruits, and fungi. Yet, the leaves of many plant species worldwide produce odour, which could act as a cue for foraging mammalian herbivores. Leaf odour may thus improve foraging efficiency for such herbivores in many ecosystems by reducing search time, particularly but not only, for plants that are visually obscured. We tested the use of leaf odour by a free-ranging mammalian browser, the swamp wallaby (Wallabia bicolor) to find and browse palatable tree seedlings (Eucalyptus pilularis). Wallabies visited patches non-randomly with respect to the presence of seedlings. In the absence of visual plant cues, they used leaf odour (cut seedlings in vials) to find patches earlier, and visited and investigated them more often than control patches (empty vials), supporting the hypothesis that wallabies used seedling odour to enhance search efficiency. In contrast, the grey kangaroo (Macropus giganteus), a grazer, showed no response to seedling odour. When the availability of seedling visual and olfactory cues was manipulated, wallabies browsed seedlings equally quickly in all treatments: upright (normal cues), pinned to the ground (reduced visual cues), and upright plus pinned seedlings (double olfactory cues). Odour cues play a critical role in food-finding by swamp wallabies, and these animals are finely tuned to detecting these cues with their threshold for detection reached by odours from only a single plant. The global significance of leaf odour in foraging by mammalian herbivores consuming conifers, eucalypts, and other odour-rich species requires greater attention.