Plant secondary metabolites alter the feeding patterns of a mammalian herbivore (<Emphasis Type="Italic">Neotoma lepida</Emphasis>)

Mammalian herbivores are predicted to regulate concentrations of ingested plant secondary metabolites (PSMs) in the blood by modifying the size and frequency of feeding bouts. It is theorized that meal size is limited by a maximum tolerable concentration of PSMs in the blood, such that meal size is predicted to decrease as PSM concentration increases. We investigated the relationship between PSM concentration in the diet and feeding patterns in the herbivorous desert woodrat (Neotoma lepida) fed diets containing phenolic resin extracted from creosote bush (Larrea tridentata). Total daily intake, meal size and feeding frequency were quantified by observing the foraging behavior of woodrats on diets containing increasing concentrations of creosote resin. Desert woodrats reduced meal size as resin concentration in the diet increased, resulting in an overall reduction in daily intake and regulation of resin intake. Moreover, desert woodrats were able to detect resin concentrations in the diet and regulate the intake of resin very rapidly. We suggest that the immediate and sustained ability to detect and regulate the intake of resin concentrations during each foraging bout provides a behavioral mechanism to regulate blood concentrations of resin and allows desert woodrats to make “wise” foraging decisions.     

Heritable variation in the foliar secondary metabolite sideroxylonal in <Emphasis Type="Italic">Eucalyptus</Emphasis> confers cross-resistance to herbivores

Plants encounter a broad range of natural enemies and defend themselves in diverse ways. The cost of defense can be reduced if a plant secondary metabolite confers resistance to multiple herbivores. However, there are few examples of positively correlated defenses in plants against herbivores of different types. We present evidence that a genetically variable chemical trait that acts as a strong antifeedant to mammalian herbivores of Eucalyptus also deters insect herbivores, suggesting a possible mechanism for cross-resistance. We provide field confirmation that sideroxylonal, an important antifeedant for mammalian herbivores, also determines patterns of damage by Christmas beetles, a specialist insect herbivore of Eucalyptus. In a genetic progeny trial of Eucalyptus tricarpa, we found significant heritabilities of sideroxylonal concentration (0.60), overall insect damage (0.34), and growth traits (0.30–0.53). Population of origin also had a strong effect on each trait. Negative phenotypic correlations were observed between sideroxylonal and damage, and between damage and growth. No relationship was observed between sideroxylonal concentration and any growth trait. Our results suggest that potential for evolution by natural selection of sideroxylonal concentrations is not strongly constrained by growth costs and that both growth and defense traits can be successfully incorporated into breeding programs for plantation trees.     

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.     

Diet switching in a generalist mammalian folivore: fundamental to maximising intake

Generalist mammalian herbivores exploit a diverse diet. A generalised feeding strategy utilises a mixed diet to obtain a range of nutrients and to reduce the detoxication load of similar groups of plant secondary metabolites (PSMs). There is limited research investigating how mammalian herbivores achieve this dietary mixing in their daily foraging activities. We investigated the patterns of, and behaviours associated with, dietary mixing in a generalist mammalian folivore, the common brushtail possum (Trichosurus vulpecula). Possums were offered foliage of two eucalypt species (Eucalyptus globulus and E. regnans) as either (a) Full choice: both species offered for 8 h; (b) Restricted choice: both species offered for 2×2 h blocks; (c) G–R no choice: E. globulus offered for the first 4 h, E. regnans offered for next 4 h; and (d) R–G no choice: E. regnans offered for first 4 h, E. globulus offered for next 4 h. We hypothesised that possums would maximise intake on the Full choice diet, where time availability was greatest in combination with a choice of foliage. We also hypothesised that diet switching, defined as the frequency of changing between food types while feeding, would play a fundamental role in maximising intake. Possums achieved maximum intake on the Full choice diet and minimum intake on the No choice diets. Although intake was similar between the Restricted choice and No choice diets, possums achieved this intake in half the amount of time when able to switch regularly between foliage on the Restricted choice diet. We conclude that a generalist herbivore’s ability to effectively switch diets when foraging is fundamental to maximising intake. Hence, the degree of plant heterogeneity in an environment, and the spatial scale at which it occurs, may affect an herbivore’s foraging decisions and, ultimately, influence its foraging efficiency.     

Behavioural contributions to the regulated intake of plant secondary metabolites in koalas

In a given period of time, herbivores often eat less as dietary plant secondary metabolite (PSM) concentrations increase. This reduction in total food intake is interpreted as a need for the herbivore to regulate PSM ingestion in order to avoid toxication. However, regulation of PSM ingestion involves more than the reduction of total intake; it involves an alteration of meal patterns, through a reduction in the number and/or the size of the meals eaten. Despite this, studies of how herbivores alter their meal patterns when offered varying concentrations of PSMs are rare. We investigated whether koalas adjust the number and/or the size of their meals when offered eucalypt foliage varying naturally in concentrations of formylated phloroglucinol compounds (FPCs), a group of PSMs that have previously been shown to inhibit total food intake. High FPC concentrations caused koalas to eat more slowly, eat shorter meals and eat less per meal, which resulted in a reduced total intake. Because increasing FPC concentrations did not cause koalas to alter the number of meals that they ate, clear individual differences between koalas were observed, where some consistently ate fewer larger meals and others ate many smaller meals. Thus, different feeding strategies may still achieve the same outcome of a regulated intake of PSMs. The changes observed match the meal patterns of other herbivores ingesting PSMs known to stimulate nausea and emetic pathways, supporting the idea that feedback signals from nausea are an important way that koalas avoid toxication when eating eucalypt foliage.     

视野受阻对东方田鼠觅食行为的影响

植食性小型哺乳动物的觅食行为是对特定生境的适应性产物。食物斑块周边植被对动物视野遮挡是否通过作用于其觅食活动中的警觉而影响摄入率。采用新鲜白三叶叶片构建东方田鼠密集均质食物斑块,以牛皮纸模拟食物斑块周边植被遮挡田鼠视野,测定其在食物斑块上的觅食行为序列过程及行为参数,检验食物斑块周边植被高度对东方田鼠摄入率的影响。结果发现,个体在不同程度视野受阻条件下食物摄入率无显著差异。分析觅食行为参数动态发现,在不同视野受阻条件下,个体能通过调整各采食回合内警觉行为动作的发生频次和持续时间,维持觅食回合内总的觅食中断时间的稳定,进而保证进食时间的稳定。东方田鼠在不同程度的视野遮挡条件下均能通过行为变异和优化使摄入率保持稳定。结果亦充分说明,东方田鼠在警觉强度上的变化不能反映觅食中断所带来的食物收益减损的代价,但觅食活动中各警觉动作的持续时间的变异却能够明确指示个体摄入率的动态变化,因此,以觅食活动中警觉引起的觅食中断时间代价为线索,检验其对摄入率的影响,是评价植食性小型哺乳动物在不同生境觅食适应性策略的有效的方法。     

Within‐plant distribution of 1,4‐benzoxazin‐3‐ones contributes to herbivore niche differentiation in maize

Plant defences vary in space and time, which may translate into specific herbivore‐foraging patterns and feeding niche differentiation. To date, little is known about the effect of secondary metabolite patterning on within‐plant herbivore foraging. We investigated how variation in the major maize secondary metabolites, 1,4‐benzoxazin‐3‐one derivatives (BXDs), affects the foraging behaviour of two leaf‐chewing herbivores. BXD levels varied substantially within plants. Older leaves had higher levels of constitutive BXDs while younger leaves were consistently more inducible. These differences were observed independently of plant age, even though the concentrations of most BXDs declined markedly in older plants. Larvae of the well‐adapted maize pest Spodoptera frugiperda preferred and grew better on young inducible leaves irrespective of plant age, while larvae of the generalist Spodoptera littoralis preferred and tended to grow better on old leaves. In BXD‐free mutants, the differences in herbivore weight gain between old and young leaves were absent for both species, and leaf preferences of S. frugiperda were attenuated. In contrast, S. littoralis foraging patterns were not affected. In summary, our study shows that plant secondary metabolites differentially affect performance and foraging of adapted and non‐adapted herbivores and thereby likely contribute to feeding niche differentiation.     

Should I stay or should I go? Patch departure decisions by herbivores at multiple scales

The question of how much time a foraging herbivore should spend in a patch of food poses a central challenge in classical foraging theory. However, there remains uncertainty about the relevance of the patch paradigm to foraging decisions by large herbivores. This paper examines evidence for successfully predicting and quantifying patch departure decisions for large mammalian herbivores foraging across several spatial and temporal scales. Departure decisions at fine scales are influenced by tradeoffs between maximizing intake rate and food quality. Classical models for departure decisions at larger spatial scales, particularly the marginal value theorem, appear inadequate. We advocate exploring alternative models for predictions of residence time at the patch scale.     

Use of trade-off theory to advance understanding of herbivore–parasite interactions

• 1 Trade‐off theory has been extensively used to further our understanding of animal behaviour. In mammalian herbivores, it has been used to advance our understanding of their reproductive, parental care and foraging strategies. Here, we detail how trade‐off theory can be applied to herbivore–parasite interactions, especially in foraging environments.
• 2 Foraging is a common mode of uptake of parasites that represent the most pervasive challenge to mammalian fitness and survival. Hosts are hypothesized to alter their foraging behaviour in the presence of parasites in three ways: (i) hosts avoid foraging in areas that are contaminated with parasites; (ii) hosts select diets that increase their resistance and resilience to parasites; and (iii) hosts select for foods with direct anti‐parasitic properties (self‐medication). We concentrate on the mammalian herbivore literature to detail the recent advances made using trade‐off frameworks to understand the mechanisms behind host–parasite interactions in relation to these three hypotheses.
• 3 In natural systems, animals often face complex foraging decisions including nutrient intake vs. predation risk, nutrient intake vs. sheltering and nutrient intake vs. parasite risk trade‐offs. A trade‐off framework is detailed that can be used to interpret mammal behaviour in complex environments, and may be used to advance the self‐medication hypothesis.
• 4 The use of trade‐off theory has advanced our understanding of the contact process between grazing mammalian hosts and their parasites transmitted via the faecal–oral route. Experimental manipulation of the costs and benefits of a nutrient intake vs. parasite risk trade‐off has shown that environmental conditions (forage quality and quantity) and the physiological state (parasitic and immune status) of a mammalian host can both affect the behavioural decisions of foraging animals.
• 5 Naturally occurring trade‐offs and the potential to manipulate their costs and benefits enables us to identify the abilities and behavioural rules used by mammals when making decisions in complex environments and thus predict animal behaviour.

     

Chemical and Mechanical Defenses Vary among Maternal Lines and Leaf Ages in Verbascum thapsus L. (Scrophulariaceae) and Reduce Palatability to a Generalist Insect

Intra-specific variation in host-plant quality affects herbivore foraging decisions and, in turn, herbivore foraging decisions mediate plant fitness. In particular, variation in defenses against herbivores, both among and within plants, shapes herbivore behavior. If variation in defenses is genetically based, it can respond to natural selection by herbivores. We quantified intra-specific variation in iridoid glycosides, trichome length, and leaf strength in common mullein (Verbascum thapsus L, Scrophulariaceae) among maternal lines within a population and among leaves within plants, and related this variation to feeding preferences of a generalist herbivore, Trichopulsia ni Hübner. We found significant variation in all three defenses among maternal lines, with T. ni preferring plants with lower investment in chemical, but not mechanical, defense. Within plants, old leaves had lower levels of all defenses than young leaves, and were strongly preferred by T. ni. Caterpillars also preferred leaves with trichomes removed to leaves with trichomes intact. Differences among maternal lines indicate that phenotypic variation in defenses likely has a genetic basis. Furthermore, these results reveal that the feeding behaviors of T. ni map onto variation in plant defense in a predictable way. This work highlights the importance of variation in host-plant quality in driving interactions between plants and their herbivores.     

饥饿和食物单宁酸对东方田鼠(Microtus fortis)食物摄入量和觅食行为的影响

实验室条件下,测定饥饿和食物单宁酸对东方田鼠食物摄入量和觅食行为的影响。结果表明,饥饿使实验个体的食物总摄入量增加,食物摄入率及口量大小随饥饿强度的增大而增加,而觅食频次则无显著改变,实验个体每取食回合的觅食时间呈缓慢增加的趋势,与对照组比较,觅食时间差异不显著。东方田鼠优先选择0%单宁酸食物,次为3%单宁酸食物,而对6%单宁酸食物的摄入量最少。在饥饿条件下,东方田鼠食物摄入率的增加主要源于其口量大小,觅食频次和觅摄食时间对食物摄入量增加的贡献不显著。在饥饿条件下,植食性小哺乳动物并未通过延长觅食时间,降低用于防卫、繁殖活动时间来增加食物摄入量,而是通过增加口量大小,提高其食物摄入率来满足其营养需要。验证了饥饿与植物次生化合物共同作用引起田鼠类动物生理的改变,能影响其食物摄入量及觅食行为的假设。     

Inter-population differences in the tolerance of a marsupial folivore to plant secondary metabolites

Plant secondary metabolites (PSMs) strongly influence diet selection by mammalian herbivores. Concentrations of PSMs vary within and among plant species, and across landscapes. Therefore, local adaptations may cause different populations of herbivores to differ in their ability to tolerate PSMs. Here, we tested the food intake responses of three populations of a marsupial folivore, the common brushtail possum (Trichosurus vulpecula Kerr), from different latitudes and habitat types, to four types of PSMs. We found clear variation in the responses of northern and southern Australian possums to PSMs. Brushtail possums from southern Australia showed marked decreases in food intake in response to all four PSMs, while the two populations from northern Australia were not as sensitive and their responses did not differ from one another. These results were unexpected, based on our understanding of the experiences of these populations with PSMs in the wild. Our results suggest that geographically separated populations of possums may have evolved differing abilities to cope with PSMs, as a result of local adaptation to their natural environments. Our results provide the basis for future studies to investigate the mechanisms by which populations of mammalian species differ in their ability to tolerate PSMs.     

Adaptive foraging of leaf‐cutter ants to spatiotemporal changes in resource availability in Neotropical savannas

1. Generalist herbivores feed on a wide and diverse set of species, but fine‐scale foraging patterns may be affected by the interplay between the quality, quantity and spatial distribution of host plants. 2. The foraging patterns of a prevalent Neotropical herbivore, the leaf‐cutter ant Atta laevigata, in the Brazilian Cerrado savannas were examined in order to determine if patterns observed are in concert with central‐place foraging predictions. 3. The results showed that A. laevigata acts as a polyphagous but highly selective herbivore, with ant attacks often resulting in partial defoliation of less‐preferred species and full defoliation of preferred ones. It was found, for the first time, that there is a strong and positive relationship between the relative attack frequency on plants from preferred species and foraging distance to the nest. This suggests a balance between the quality of plant resources harvested and costs involved in their transportation. It was also observed that colonies focused their harvest on preferred species in months with low availability of young leaves. Consequently, high herbivory rate was more frequent in plants attacked far away from the nest and in dry months. 4. These assessments highlight the fact that Atta colonies may become more selective as foraging distance to the nest increases and in response to fluctuations in the availability of palatable resources throughout the year. The results also show some dissimilarities in the foraging behaviours of A. laevigata when compared with other locations, suggesting that widely distributed herbivores may modify foraging strategies across their geographic range.     

Evolutionary irony: evidence that ‘defensive’ plant spines act as a proximate cue to attract a mammalian herbivore

Many plants produce structural defenses to deter feeding by herbivores. However, many previous studies testing whether spines are effective at defending against mammalian herbivores have produced equivocal results. These ambiguous results are hypothesized to be due to herbivore counter‐adaptations. We investigated potential counter‐adaptations in a population of white‐throated woodrats Neotoma albigua that specialize on cactus by investigating feeding behavior and preference for cacti varying in spinescence. Neotoma albigula exhibited a unique behavior of clipping cactus spines, which renders these defenses ineffective. Strikingly, these woodrats chose to collect spiny cacti over experimentally de‐spined cacti, demonstrating that spines act as a proximal cue that attracts woodrats. This attraction is likely due to the higher protein and lower fiber content of spiny cacti compared to naturally non‐spiny cacti. Thus, the ‘defensive’ spines of cacti are ineffective against a specialist herbivore and instead serve as an indicator of nutritional quality that promotes herbivory. Our results support the ‘rule‐of‐thumb’ hypothesis of foraging, which states that herbivores forage according to obvious visual cues that are indicative of nutritional content, rather than sampling nutrient composition of plants. We propose that specialist herbivores are unique systems in which to study other counter‐adaptations to structural defenses and ‘rule‐of‐thumb’ foraging strategies.     

Drought changes plant chemistry and causes contrasting responses in lepidopteran herbivores

Drought events are predicted to increase due to climate change, yet consequences for plant–insect interactions are only partially understood. Drought‐mediated interactions between herbivores and their host plants are affected by a combination of factors, including characteristics of the affected plant, its associated herbivore and of the prevailing drought. Studying the effect of these factors in combination may provide important insight into plant and herbivore responses to drought. We studied drought effects on plant resistance to two leaf‐chewing herbivores by considering differing growth conditions, plant chemistry and insect responses in concert. We exposed Alliaria petiolata plants from several wild populations to different intensities of intermittent drought stress and quantified drought‐mediated changes in plant chemistry. Simultaneously, we assessed behavior (feeding preference) and performance of two lepidopteran herbivores: Pieris brassicae, a specialist, and Spodoptera littoralis, a generalist. Drought led to lowest concentrations of secondary defense compounds in severely stressed plants, without affecting total nitrogen content. Additionally, drought evoked opposite patterns in feeding preferences (plant palatability) between the herbivore species. Pieris brassicae consumed most of well‐watered plants, while S. littoralis preferred severely drought‐stressed plants. Hence, feeding preferences of S. littoralis reflected changes in plant secondary chemistry. Contrary to their feeding preference, P. brassicae performed better on drought‐stressed than on well‐watered plants, with faster development and higher attained pupal mass (plant suitability). Spodoptera littoralis showed retarded development in all treatments. In conclusion, drought caused plant secondary defense compounds to decrease consistently across all studied plant populations, which evoked contrasting feeding preferences of two herbivore species of the same feeding guild. These results suggest herbivore specificity as a possible explanation for herbivore responses to drought and emphasize the importance of herbivore characteristics such as feeding specialization in understanding and predicting consequences of future drought events.     

Herbivore species identity rather than diversity of the non‐host community determines foraging behaviour of the parasitoid wasp Cotesia glomerata

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species‐rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore‐induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non‐suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non‐host herbivores influence the behaviour of parasitoids, we created environments with different levels of non‐host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non‐host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non‐host species diversity nor non‐host identity influenced the preference of the parasitoid for herbivore‐infested plants. However, after landing on the plant, non‐host species identity did affect parasitoid behaviour, whereas non‐host diversity did not. One of the non‐host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non‐host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non‐host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.     

What physiological processes permit insects to eat Eucalyptus leaves?

Many species of insects eat Eucalyptus foliage despite its relatively low nutritional value and the many plant secondary metabolites (PSMs) present, for example, terpenes, phenols and formylated phloroglucinols (FPGs). Formylated phloroglucinols are a new class of PSMs that act as antifeedants for possums and koalas. What physiological processes are present that permit insects to eat eucalypt foliage and how do PSMs influence insect feeding or digestion? Some trees seem to be repeatedly infested with eucalypt‐feeding insects, possibly as a result of previous chemosensory cues remaining from parental selection of a plant. Avoidance or storage of PSMs permit jarrah leafminers (Perthida glyphopa) and sawflies (Perga sp.) to consume eucalypt foliage without dealing with the majority of these compounds. Some PSMs can be metabolized by polysubstrate membrane oxidases as found in caterpillars or sawflies that feed on eucalypts. High midgut pH may be advantageous for nutrient extraction and PSM metabolism, and midgut pH ranges between 8.5 and 8.9 for caterpillars of Hyalarcta huebneri. Plant secondary metabolites may not be absorbed as a result of the combined presence of the peritrophic matrix and endogenous surfactants. Excretion of PSMs can be as metabolites or intact compounds. Both putative metabolites and sideroxylonal‐A, an FPG, are present in the faeces of larvae of the case moth, H. huebneri. The presence of sideroxylonal‐A in the food had an effect on the presence of 5‐hydroxytryptamine (5HT) in the central nervous system of caterpillars, as larvae fed leaves with a high concentration of sideroxylonal‐A had relatively more 5HT in the brain and central nervous system ganglia than larvae fed leaves containing a low concentration. Further work is necessary to clarify how PSMs are handled by eucalypt‐feeding insects and what effect FPGs have on feeding and digestion.     

Foraging behaviour at multiple temporal scales in a wild alpine equid

Forage abundance, forage quality, and social factors are key elements of the foraging ecology of wild herbivores. For non-ruminant equids, forage-limited environments are likely to impose severe constraints on their foraging behaviour. We used a multi-scale approach to study foraging behaviour in kiang (Equus kiang), a wild equid inhabiting the high-altitude rangelands of the Tibetan Plateau. Using behavioural observations and vegetation sampling, we first assessed how patterns of plant abundance and quality affected (i) the instantaneous forage intake rate (fine scale) and (ii) the proportion of time spent foraging (coarse scale) across seasons. We also tested whether foraging behaviour differed among group types, between sex in adults, and between females of different reproductive status. At a fine scale, intake rate increased linearly with bite size and increased following a type II curvilinear function with biomass on feeding sites. Forage intake rate also increased linearly with plant quality. Male and female kiangs had similar intake rates. Likewise, gravid and lactating females had similar intake rates as barren and non-lactating females. At a coarse scale, kiangs spent longer time feeding in mesic than in xeric habitats, and spent more time feeding in early summer and fall than in late summer. Groups of adults with foals spent less time feeding than male groups and groups of adults without foals. Our findings suggest that kiangs use flexible foraging behaviours in relation to seasonal variations of vegetation quality and abundance, a likely outcome of the extreme seasonal conditions encountered on the Tibetan Plateau.     

Fatal attraction: vegetation responses to nutrient inputs attract herbivores to infectious anthrax carcass sites

Parasites can shape the foraging behaviour of their hosts through cues indicating risk of infection. When cues for risk co-occur with desired traits such as forage quality, individuals face a trade-off between nutrient acquisition and parasite exposure. We evaluated how this trade-off may influence disease transmission in a 3-year experimental study of anthrax in a guild of mammalian herbivores in Etosha National Park, Namibia. At plains zebra (Equus quagga) carcass sites we assessed (i) carcass nutrient effects on soils and grasses, (ii) concentrations of Bacillus anthracis (BA) on grasses and in soils, and (iii) herbivore grazing behaviour, compared with control sites, using motion-sensing camera traps. We found that carcass-mediated nutrient pulses improved soil and vegetation, and that BA is found on grasses up to 2 years after death. Host foraging responses to carcass sites shifted from avoidance to attraction, and ultimately to no preference, with the strength and duration of these behavioural responses varying among herbivore species. Our results demonstrate that animal carcasses alter the environment and attract grazing hosts to parasite aggregations. This attraction may enhance transmission rates, suggesting that hosts are limited in their ability to trade off nutrient intake with parasite avoidance when relying on indirect cues.     

The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions

Diet selection by mammalian herbivores is often influenced by plant community composition, and numerous studies have focused on the relationships between herbivore foraging decisions and food/plant species abundance. However, few have examined the role of neighbour palatability in affecting foraging of a target plant by large mammalian herbivores. We used a large-scale field dataset on diet selection by red deer Cervus elaphus in Fiordland National Park, New Zealand to: (1) estimate the palatability of native forest plant species to introduced deer from observed patterns of browse damage; and (2) examine whether intraspecific variation in browsing of plants can be related to variation in the local abundance of alternative forage species. Overall, 21 of the 53 forest species in our dataset were never browsed by deer. At a community level, plants were more likely to be browsed if they were in a patch of vegetation of high forage quality, containing high abundances of highly palatable species and/or low abundances of less-palatable species. Our findings suggest that deer make foraging decisions at both a coarse-grain level, selecting vegetation patches within a landscape based on the overall patch quality, and at a fine-grain level by choosing among individual plants of different species.