Mammalian Browsers Research Articles

Dynamic effects of insect herbivory and climate on tundra shrub growth: Roles of browsing and ramet age

Abstract To predict shrub responses under climate change in tundra, we need to understand how thermal conditions and herbivory contribute to growth. We hypothesise that shrub growth increases with thermal conditions and precipitation, but that this increase is counteracted by insect herbivory, and that these climate–insect herbivory relationships are modified by both browsing and plant age. We use empirical dynamic modelling (EDM) to analyse a 20‐year time series on willow Salix phylicifolia shoot growth, growing degree days, summer precipitation and herbivory from an experiment at forest–tundra ecotone. The experiment includes manipulations of avian and mammal browsing (fences) and ramet age (pruning to rejuvenate willows). Negative effects of insect herbivory on willow shoot growth were intensified during warmer years, whereas increasing precipitation led to reduced effects. Moreover, the effect of insect herbivores on shoot growth varied with ramet age and vertebrate browsing: younger ramets generally experienced less negative insect herbivore effects, whereas ptarmigan browsing was associated with more positive temperature effects on shoot growth, and reindeer browsing with more negative effects of insect herbivory and precipitation. Synthesis. Our findings show that the negative effects of insect herbivory on shoot growth likely intensify under warmer thermal conditions, but that increasing precipitation can counteract these effects. Moreover, changes in thermal conditions, precipitation and vertebrate browsers all have predictable, albeit complex and nonlinear, effects on shrub growth, highlighting the importance of long‐term experimental data and flexible analytical methods such as EDM for characterising climate and community interactions in artic systems.

The role of browsers in maintaining the openness of savanna grazing lawns

Abstract In savannas, ruminant herbivores can have divergent impacts on tree recruitment and subsequent woody cover. Whereas heavy grazing by cattle results in woody thickening, intensive grazing by wildlife instead tends to be associated with lower woody cover. To disentangle why woody cover is low in areas heavily grazed by wildlife, we investigated (a) whether grazing lawns attract indigenous mammalian browsers, and if a preference for short‐grass habitat decreases with browser body mass as predator susceptibility decreases; and (b) whether browser attraction to grazing lawns translates into the suppression of woody plants, including seedlings and saplings, thus maintaining the openness of heavily grazed short‐grass areas. In Kruger National Park, South Africa, we contrasted browser abundance (using dung counts) on grazing lawns and on low‐herbivory sites characterised by tall grass. Additionally, a herbivore exclosure experiment was set up to investigate the combined impact of browser removal and grass height habitat type on seedling survival and sapling growth of a dominant woody plant species. Finally, in Hluhluwe‐iMfolozi Park (HiP), we examined the long‐term (10 years) impact of browser removal on the growth rates of a range of woody species, monitored across ten sites along a gradient of herbivory ranging from grazing lawn to tall grass. Steenbok and impala selected short‐ over tall grass as preferred browsing sites, while elephant preferred tall grass. Browser abundance on short grass decreased with browser body mass, indicating that predator avoidance might be a key factor driving mesoherbivores to utilise grazing lawns. Seedling survival was lowest on grazing lawns when browsers were present, with mortality occurring in two out of every three seedlings. Similarly, sapling growth was lowest on grazing lawns, although browser removal had no significant effect. Evidence for increased browser impact on grazing lawns was clearest from our long‐term herbivore exclosure experiment in HiP, which demonstrated that browsers strongly modify the growth rates of woody plants in short‐grass habitats. Synthesis. These results provide support for the hypothesis that browsers, particularly browsing mesoherbivores and mixed feeders, are attracted to short‐grass habitats, and that they help maintain grazing lawn openness by suppressing seedling survival and woody plant growth where grass is kept short by grazers.

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.

Acacia Ants Respond to Plant-Borne Vibrations Caused by Mammalian Browsers

Living in the African savanna is dangerous, especially for plants. Many plants therefore engage in mutualism with ants, in which plants provide food and shelter in exchange for protection against herbivores. Ants become alarmed when the plant takes on some sort of damage. They immediately emerge from their plant shelter and aggressively defend the plant. Mammalian herbivores can have devastating effects on trees by browsing, breaking tree branches, stripping bark, and pushing over entire trees. However, mutualistic ants substantially reduce the amount of damage. To efficiently protect the tree, ants need to rapidly react together when the tree is under attack. Here, we show that the acacia ant Crematogaster mimosae defends its host tree by exploiting plant-borne vibrations caused by browsers feeding on the tree. Experiments with controlled vibrations show that ants discriminate browser-induced vibrations from those induced by wind, become alarmed, and patrol on the branches. Browser-induced vibrations serve as a long-distance alarm cue. The vibrations propagate through the whole acacia tree and trigger ants' defensive behavior, even on the other side of the tree. Furthermore, the ants make use of tropotactic directional vibration sensing to orient to the attacked part of the tree and fight back the attacker.

Heterospecific Tree Density and Environmental Factors Affect Afzelia africana Sm. Population Structure in the Pendjari Biosphere Reserve, West Africa: Implications for Management and Restoration

Information on how abiotic and biotic factors affect species population structures and regeneration are critical for understanding plant growth in natural habitats. Here, we used the data from three spatially distinct populations of Afzelia africana Sm. in the Pendjari Biosphere Reserve in Benin, to determine how the species population structures respond to abiotic and biotic factors. Afzelia africana population structures were studied using several parameters including basal area, tree height, density of successive diameter classes, and size class slope. We tested for individual effects of abiotic (mound density, soil type, and terrain slope) and biotic (heterospecific tree density) factors on the species population structure. We also tested for similarity of species composition among studied A. africana population stands. Results revealed a tree density structure with mature individuals, and size class distribution indicating a recruitment bottleneck at the juvenile stage (10–20 cm diameter), possibly due to mammal browsing, natural and artificial fires. Heterospecific tree density was positively associated with A. africana adult density but negatively related to the species growth parameters (mean diameter, basal area, and tree height). These results indicate some degrees of niche overlap between A. africana and coexisting species but also partly reflect A. africana tolerance and adaptation to limited resources environment. Soil type significantly influenced both basal area and regeneration density, greater values being observed on silt-sand-rocky soils. Basal area was higher on steeper slope, probably a result of species conservative strategies. These findings were discussed in line with management and restoration action needs in the Pendjari Biosphere Reserve.

Leaf herbivory by insects during summer reduces overwinter browsing by moose

BackgroundDamage to plants by herbivores potentially affects the quality and quantity of the plant tissue available to other herbivore taxa that utilize the same host plants at a later time. This study addresses the indirect effects of insect herbivores on mammalian browsers, a particularly poorly-understood class of interactions. Working in the Alaskan boreal forest, we investigated the indirect effects of insect damage to Salix interior leaves during the growing season on the consumption of browse by moose during winter, and on quantity and quality of browse production.ResultsTreatment with insecticide reduced leaf mining damage by the willow leaf blotch miner, Micrurapteryx salicifoliella, and increased both the biomass and proportion of the total production of woody tissue browsed by moose. Salix interior plants with experimentally-reduced insect damage produced significantly more stem biomass than controls, but did not differ in stem quality as indicated by nitrogen concentration or protein precipitation capacity, an assay of the protein-binding activity of tannins.ConclusionsInsect herbivory on Salix, including the outbreak herbivore M. salicifoliella, affected the feeding behavior of moose. The results demonstrate that even moderate levels of leaf damage by insects can have surprisingly strong impacts on stem production and influence the foraging behavior of distantly related taxa browsing on woody tissue months after leaves have dropped.

A lepidopteran (Imbrasia belina) might influence tree-grass balance of Colophospermum mopane savanna

Traditional explanations of tree-grass coexistence in African savannas are based on competition between these growth forms or demographic bottlenecks of trees maintained by fire or mammalian browsers. Perturbation of their “balance” may result in an alternate system state of woody encroachment. Invertebrate herbivory has never been offered as an explanation. We developed a consumer-resource model which illustrated that annual irruptions of a lepidopteran (Imbrasia belina), known as mopane worm, can determine the tree-grass balance of semi-arid Colophospermum mopane savanna in southern Africa. Model performance was sensitive to the abundance, hence mortality, of mopane worms, owing to their complete defoliation of tree leaf biomass resulting in altered competitive relations between trees and grasses. Invertebrate herbivores have been recognized in other systems as agents for effecting a state change of host tree populations; this modeling study offers a first indication of such a role for the well-researched tree-grass relations of African savannas.

Induction and persistence of allelochemicals in the foliage of balsam fir seedlings following simulated browsing

The intensity, pattern, and timing of browsing disturbances influence the mobilization of chemical defenses (allelochemicals) and subsequent growth of conifers such as balsam fir, an important food source for various insect and mammalian herbivores. The objective of this study was to examine the induction and persistence of allelochemicals in foliage of balsam fir seedlings as affected by the pattern and timing of browsing disturbance. We conducted a simulated browsing study in a greenhouse environment using four browsing patterns and three persistence times. Total phenols were induced shortly after simulated browsing and persisted for at least 1 month, whereas condensed tannins exhibited a delayed induction but were more persistent, remaining above background levels 2 months after simulated browsing. The chemistry of non-browsed seedlings revealed that a seasonal pattern was evident for both total phenols and condensed tannins. As the experiment progressed through the growing season, background levels of total phenols decreased while those of condensed tannins increased with the two allelochemicals groups appearing to show a negative relationship. These trends suggest that total phenols, which are assumed to have a lower biosynthetic cost and are more quickly mobilized than condensed tannins, are a first line of herbivory defense followed by condensed tannins which take longer to mobilize but have a longer persistence time or that total phenols act as a primary defense against insect herbivores which disturb plants in the early growing season while condensed tannins are mobilized in the late season to defend against mammalian browsers of balsam fir in winter.

Borrowed plant defences: Deterring browsers using a forestry by-product

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.

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.

Follow your nose: leaf odour as an important foraging cue for mammalian herbivores.

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.

Accounting for the nested nature of genetic variation across levels of organization improves our understanding of biodiversity and community ecology

Recent work has demonstrated that the presence or abundance of specific genotypes, populations, species and phylogenetic clades may influence community and ecosystem properties such as resilience or productivity. Many ecological studies, however, use simple linear models to test for such relationships, including species identity as the predictor variable and some measured trait or function as the response variable without accounting for the nestedness of genetic variation across levels of organization. This omission may lead to incorrect inference about which source of variation influences community and ecosystem properties. Here, we explicitly compare this common approach to alternative ways of modeling variation in trait data, using simulated trait data and empirical results of common‐garden trials using multiple levels of genetic variation within Eucalyptus, Populus and Picea. We show that: 1) when nested variation is ignored, an incorrect conclusion of species effect is drawn in up to 20% of cases; 2) overestimation of the species effect increases – up to 60% in some scenarios – as the nested term explains more of the variation; and 3) the sample sizes needed to overcome these potential problems associated with aggregating nested hierarchical variation may be impractically large. In common‐garden trials, incorporating nested models increased explanatory power twofold for mammal browsing rate in Eucalyptus, threefold for leaf area in Populus, and tenfold for branch number in Picea. Thoroughly measuring intraspecific variation and characterizing hierarchical genetic variation beyond the species level has implications for developing more robust theory in community ecology, managing invaded natural systems, and improving inference in biodiversity–ecosystem functioning research.SynthesisUntil recently, ecologists acknowledged the ubiquity of within‐species trait variation, but paid scant attention to how much it affects communities and ecosystems. Here, the authors used simulated trait data and common‐garden studies to demonstrate that we ignore intraspecific trait variation at our peril. In both simulated and experimental systems, in many cases ignoring intraspecific variation led to incorrect statistical inferences and inflated the effect size of species identity.This study shows that ecologists must characterize hierarchically nested genetic and phenotypic variation to fully understand the links between individual traits, community structure and ecosystem functioning.

Recruitment of a keystone tree species must concurrently manage flooding and browsing

Summary Multiple pressures (land‐use change, water extraction and climate change) interact to influence biodiversity and ecosystem processes, but direct evidence for interactions among multiple pressures is limited. Floodplain forests are an acute example of how interacting pressures (river regulation, water extraction, decreasing rainfall and mammal browsing) interact to degrade native ecosystems. We conducted a 2‐year field experiment to determine how flooding, browsing and sediment salinity interacted to determine in situ seedling survival and growth of the keystone floodplain tree species (Eucalyptus camaldulensis Dehnh.). On semi‐arid floodplains of southern Australia, 1‐year‐old seedlings were planted on the banks of six ephemeral creeks, three of which were flooded with management flows before planting while the others remained dry. Four plots were established at each creek, two open to browsing and two fenced to exclude mammal herbivores. Flooding had a strong positive effect on seedling survival and height, but browsing had strong negative effects. Sediment salinity (a covariate rather than a designed effect) had a weak negative effect on seedling survival and height. The positive effects of flooding were largely offset by the negative interaction with browsing and, to a lesser extent, sediment salinity. Although flooding has been restored to some degraded floodplain forests subjected to river regulation and a drying climate, the long‐term success of such actions is likely to be undermined by persistent browsing. Synthesis and applications. Management actions that focus on single pressures (e.g. infrequent flooding) and processes (e.g. mature tree survival) while ignoring other pressures are unlikely to sustain populations of keystone species, suggesting that complementary strategies (managed flooding with herbivore control) are necessary to sustain recruitment and, therefore, ensure the future health of these essential ecosystems.

An index for assessing effectiveness of plant structural defences against mammal browsing

Plant structural defences against mammals play an important role in ecosystem functioning as they simultaneously mediate the fitness of both animals and plants. The efficiency of structural defences can be described by the amount of plant material an animal can remove in one bite. Quantifying bite size by direct observation is difficult requiring controlled access to both the plants of interest and the herbivores. A method called the bite size index (BSI) has been proposed using human bites to simulate the bite size of medium-sized mammalian herbivores. In this paper, we evaluated the utility of the BSI for assessing the efficacy of plant structural defences. We analysed the intra- and inter-specific variability of the BSI in an African savanna (Hluhluwe–iMfolozi, South Africa) rich in herbivores, its repeatability between recorders, how the results reflect known properties of plant structural defences and how well the index correlates with the bite size of goats and its efficacy for explaining species distribution on a fire–herbivory gradient. The BSI was readily applied to a large set (n = 55) of plant species and proved to be consistent among recorders (8 of the 9 recorders obtained similar estimates). Bite sizes of goats and human observers were strongly correlated for 14 plant species (R2 = 0.70) with divergent leaf and branch types. An analysis of BSI for 55 plant species disclosed two main contributory factors: leaf dimensions and spinescence. The BSI was significantly related to plant species distribution on a fire–herbivore gradient in an African savanna.

No release for the wicked: enemy release is dynamic and not associated with invasiveness.

The enemy release hypothesis predicts that invasive species will receive less damage from enemies, compared to co-occurring native and noninvasive exotic species in their introduced range. However, release operating early in invasion could be lost over time and with increased range size as introduced species acquire new enemies. We used three years of data, from 61 plant species planted into common gardens, to determine whether (1) invasive, noninvasive exotic, and native species experience differential damage from insect herbivores. and mammalian browsers, and (2) enemy release is lost with increased residence time and geographic spread in the introduced range. We find no evidence suggesting enemy release is a general mechanism contributing to invasiveness in this region. Invasive species received the most insect herbivory, and damage increased with longer residence times and larger range sizes at three spatial scales. Our results show that invasive and exotic species fail to escape enemies, particularly over longer temporal and larger spatial scales.

Herbivore search behaviour drives associational plant refuge

Herbivore foraging decisions leading to consumption of a plant are complex and multi-faceted, shaped both by the plant itself and by its neighbours. Associational plant refuge arises when neighbours reduce focal plant susceptibility to herbivory. The specific foraging behaviours generating refuge patterns have rarely been examined in free-ranging systems, yet these are key to understanding why such refuge works or fails. We aimed to integrate herbivore foraging and associational plant refuge theories by linking foraging decisions directly to browsing outcomes on focal plants and their neighbours. We tested whether obstructive, unpalatable neighbours reduce the number of patch visits and/or interrupt searching, leading to associational refuge of focal plants. We compared visits by and behaviours of free-ranging mammalian browsers, swamp wallabies (Wallabia bicolor), in control and manipulated plant patches using cameras. Patches (7 m2) comprised a central focal plant (palatable native tree seedling, Eucalyptus pilularis) with neighbours of either existing or manipulated vegetation (unpalatable native daisy, Coronidium elatum). Wallabies made fewer visits to control than manipulated patches, but always browsed the focal plant during the first visit to a control patch. In contrast, wallabies often visited manipulated patches multiple times before browsing the focal plant. These ‘futile’ visits were both shorter and involved less searching time than visits when the focal plant was browsed. Focal plants escaped browsing for longer in manipulated than in control patches, and although none had escaped browsing after one year, survival was significantly greater in manipulated patches. We demonstrate that reduced investment in searching during visits to manipulated patches drove the associational plant refuge, but this refuge was eventually surmountable. Understanding the behaviours underpinning refuges allows better prediction of outcomes, and explains why refuge can collapse. By shaping foraging behaviour in patches, neighbouring vegetation can increase the probability that palatable plants persist despite high herbivore pressure.

Century‐scale effects of invasive deer and rodents on the dynamics of forests growing on soils of contrasting fertility

Understanding the long‐term impacts of invasive mammalian browsers and granivores in mixed forests is difficult due to the many processes potentially affecting the demography of long‐lived trees. We constructed individual‐based spatially explicit simulation models of two mixed conifer–angiosperm forests, growing on soils of contrasting phosphorus (P) availability, to investigate how browsing by invasive red deer (Cervus elaphus scoticus) and granivory by invasive rodents (primarily house mouse Mus musculus) might alter forest dynamics. Models were parameterized with field data. Seedling growth and survival rates were estimated inside and outside deer exclosures. Seed predation rates were estimated at high and low rodent densities in mast and non‐mast seeding years. For the alluvial terrace forest, which grew on P‐rich soil, our model contained 15 tree species dominated by angiosperms; our model of the P‐poor marine terrace forest contained seven species dominated by conifers. The two forest models were used to explore four 500‐year scenarios: deer and rodents present, deer present and rodents absent, deer absent and rodents present, and deer and rodents absent.Our field studies revealed that the highest probabilities of seed predation by rodents occurred for two canopy species in mast years and in neighborhoods of high species‐specific basal area, and that deer browsing had the greatest negative effects on the growth and survival rates of angiosperm seedlings. Our simulation models predicted that the presence of deer and rodents would reduce the abundances of canopy codominants, and hence total basal area, in the alluvial terrace forest. The presence of deer increased the dominance of conifers in both forests, but effects of deer and rodents were much stronger in the alluvial terrace forest. Our study revealed two emergent properties. First, rodent‐ and deer‐induced reductions in seedling abundances did not always translate into reduced sapling and adult tree abundances. Second, when deer changed the abundance of a canopy dominant, other species were affected by altered interspecific competition.We conclude that the effects of invasive deer and rodents will be greatest in forests growing on P‐rich soils. Exclusion of deer is predicted to have greater long‐term effects on forest dynamics than exclusion of rodents.

Feeding rates of a mammalian browser confirm the predictions of a ‘foodscape’ model of its habitat

Adequate nutrition is a fundamental requirement for the maintenance and growth of populations, but complex interactions between nutrients and plant toxins make it difficult to link variation in plant quality to the ecology of wild herbivores. We asked whether a 'foodscape' model of habitat that uses near-infrared spectroscopy to describe the palatability of individual trees in the landscape, predicted the foraging decisions of a mammalian browser, the koala (Phascolarctos cinereus). Specifically, we considered four behavioural decision points at which nutritional quality may influence an animal's decision. These were: which tree to enter, whether to feed from that tree, when to stop eating, and how long to remain in that tree. There were trends for koalas to feed in eucalypt trees that were more palatable than unvisited neighbouring conspecific trees, and than trees that they visited but did not eat. Koalas ate longer meals in more palatable trees, and stayed longer and spent more time feeding per visit to these trees. Using more traditional chemical analyses, we identified that an interaction between the concentrations of formylated phloroglucinol compounds (a group of plant secondary metabolites) and available N (an integrated measure of tannins, digestibility and N) influenced feeding. The study shows that foodscape models that combine spatial information with integrated measures of food quality are a powerful tool to predict the feeding behaviour of herbivores in a landscape.

Translating nutritional ecology from the laboratory to the field: milestones in linking plant chemistry to population regulation in mammalian browsers

A central goal of nutritional ecology is to understand how variation in food quality limits the persistence of wild animal populations. Habitat suitability for browsing mammals is strongly affected by concentrations of nutrients and plant secondary metabolites (PSMs), but our understanding of this is based mostly on short‐term experiments of diet selection involving captive animals. In the wild, browsers forage in biologically, chemically and spatially‐complex environments, and foraging decisions in response to varying food quality will be correspondingly complicated. We have identified four steps that must be achieved in order to translate our understanding from laboratory experiments to populations of mammalian browsers: 1) knowing what foods and how much of these wild browsers eat, as well as what they avoid eating; 2) knowing the relevant aspects of plant nutritional and defensive chemistry to measure in a given system and how to measure them; 3) understanding the spatial distribution of nutrients and PSMs in plant communities, the costs they impose on foraging and the effects on animals’ distributions; and 4) having appropriate statistical tools to analyse the data. We discuss prospects for each of these prerequisites for extending laboratory studies of nutritional quality, and review recent developments that may offer solutions for field studies. We also provide a synthesis of how to use this nutritional knowledge to link food quality to population regulation in wild mammals and describe examples that have successfully achieved this aim.

Effects of tree species richness and composition on moose winter browsing damage and foraging selectivity: an experimental study

The optimal foraging theory, the nutrient balance hypothesis, and the plant association theories predict that foraging decisions and resulting tree damage by large mammalian browsers may be influenced by the species richness and species composition of forest stands. This may lead to either associational susceptibility (increased damage on a focal plant in a mixed stand) or associational resistance (reduced damage in a mixed stand). Better understanding of the mechanisms and the relative importance of tree species richness and composition effects on foraging by mammalian browsers is needed to support sustainable management of forests and mammal populations. However, existing knowledge of forest diversity effects on foraging by large mammalian browsers comes largely from observational studies while experimental evidence is limited. We analysed winter browsing by moose (Alces alces L.) in a long-term, large-scale experiment in Finland, which represents a tree species richness gradient from monocultures to 2-, 3- and 5-species mixtures composed of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.), Siberian larch (Larix sibirica Ledeb.), silver birch (Betula pendula Roth.) and black alder (Alnus glutinosa L.). The intensity of browsing per plot increased with tree species richness while browsing selectivity decreased with tree species being targeted more equally in species-rich mixtures. Tree species composition of a plot was also an important determinant of intensity of browsing. The greatest browsing occurred in plots containing preferred species (pine and birch) while intermediate preference species (larch and alder) experienced associational susceptibility when growing with pine and birch compared with their monocultures or mixtures without pine and birch. In contrast, we found no evidence of associational resistance; the presence of a least preferred species (spruce) in a mixture had no significant effect on moose browsing on other tree species. We demonstrate that the presence of alternative forage species allows moose to spend longer opportunistically foraging in a plot, resulting in increased level of damage in species-rich stands and stands containing preferred tree species. Our results highlight the limitations of the optimal foraging theory in predicting browsing patterns and demonstrate the importance of associational effects within mixed stands.