Optimal Diet Theory Research Articles

A combination of rules govern fruit trait preference by frugivorous bat and bird species: nutrients, defence and size

Several hypotheses explain how sensory, anatomical and physiological constraints drive fruit preference in frugivores. Optimal diet theory (ODT) states that frugivores make decisions based on the energy contained in food. In contrast, geometry of nutrition (GN) states that animals balance their macronutrient intake instead, opting for rough energy. The defence trade-off hypothesis (DTH) assumes a negative relationship between secondary compounds and fruit preference. Finally, the size-matching hypothesis (SMH) states that frugivores are more attracted to fruits that are easier to handle and consume. We tested these four hypotheses by offering paired fruit species to three fruit-eating animal species in captivity that either chew fruit in the beak or mouth (‘masher’: 1 bird species, 1 bat species) or swallow whole fruits (‘gulper’: 1 bird species), from which we built a ranking of fruit preference. We then explored the importance of 13 fruit traits in explaining fruit preference. The masher bird was the only species whose fruit preference pattern corresponded with GN. Fruit preference of the masher and gulper bird species supported DTH and SMH, while fruit preference by the bat species was not related to any analysed trait. More than one single rule governs fruit preference in different frugivore species. Fruit preferences of functionally different frugivore species are affected by particular fruit traits, which they either select or avoid. The search for specific macronutrients, while avoiding toxicity in fruits matching the anatomical limitations of frugivores, could lead to complementary foraging choices. Variability between seed disperser species in their search for easily manageable fruits allows achieving a complementary fruit diet, with preferred and avoided traits. It can be an important driver of fruit trait diversity in fleshy-fruited plant assemblages. • Fruit macronutrient content and secondary compounds influence frugivore preferences. • Fruit size also influences frugivores' ability to interact with fruits. • Frugivore species differ in their search strategies to obtain complementary diets. • Preferred and avoided traits can drive diversity of fleshy-fruited plant assemblages.

Increasing availability of palatable prey induces predator-dependence and increases predation on unpalatable prey

Understanding the factors governing predation remains a top priority in ecology. Using a dragonfly nymph-tadpole system, we experimentally varied predator density, prey density, and prey species ratio to investigate: (i) whether predator interference varies between prey types that differ in palatability, (ii) whether adding alternate prey influences the magnitude of predator interference, and (iii) whether patterns of prey selection vary according to the predictions of optimal diet theory. In single-prey foraging trials, predation of palatable leopard frog tadpoles was limited by prey availability and predator interference, whereas predation of unpalatable toad tadpoles was limited by handling time. Adding unpalatable prey did not affect the predator’s kill rate of palatable prey, but the presence of palatable prey increased the influence of predator density on the kill rate of unpalatable prey and reduced unpalatable prey handling time. Prey selection did not change with shifts in the relative abundance of prey types. Instead, predators selected easy-to-capture unpalatable prey at low total densities and harder-to-capture palatable prey at high densities. These results improve our understanding of generalist predation in communities with mobile prey, and illustrate that characteristics of the prey types involved govern the extent to which alternate prey influence the predator’s kill rate.

Weighted individual-resource networks in prey-predator systems: the role of prey availability on the emergence of modular structures.

Ecological networks, usually depicting interactions among species, have been recently down-scaled to the individual level, permitting description of patterns of inter-individual resource variation that are usually hindered at the species level. Optimal diet theory (ODT) models, applied to prey-predator systems, predict different patterns of nestedness and modularity in the network, depending on the available resources and intra-specific competition. The effect of resource availability on the emergence of networks structures, and ODT framework, has not yet fully been clarified. Here, we analyzed the structural patterns of individual-resource networks in 3 species of Mediterranean salamanders, in relation to changes in prey availability. We used weighted individual-resource network metrics to interpret the observed patterns, according to 3 ODT models. We found significant nestedness recurring in our study system, indicating that both selective and opportunistic individuals occur in the same population. Prey diversity, rather than abundance, was apparently related to inter-individual resource variation and promoted the emergence of significant modularity within all networks. The observed patterns of nestedness and modularity, together with the variation in resource diversity and intra-specific competition, are in agreement with the distinct preferences model of ODT. These findings suggest that in the focal prey-predator systems, individuals were able to perceive changes in prey diversity and to exploit in different ways the variations in composition of available resources, shifting their diet assembly rules accordingly. Our findings also confirm that the use of weighted individual-resource networks, in prey-predator systems, allows to disclose dynamics that are masked at the species or population level.

Blue mussel (Mytilus edulis L.) association with conspecifics affects mussel size selection by the common seastar (Asterias rubens L.)

Prey selection by predators is of interest to community ecologists. By choosing some prey over others, predators affect prey population dynamics and the strength of food webs. According to Optimal Diet Theory (ODT), as prey density increases predators will select more profitable prey. Thereby, prey population dynamics can also affect predator behaviour. Prey profitability involves the prey energetic value, energetic costs associated to predation (time spent searching, capturing and handling prey) and certain characteristics such as body size, abundance and behaviour. We examined the effect of the blue mussel (Mytilus edulis) association with conspecifics at different densities on prey selection and behaviour by the seastar Asterias rubens. Contrary to ODT prediction, when mussels were tightly clumped in high densities, the size range of the mussels consumed by A. rubens did not differ of the size range of the mussel population. Mussel association at high density caused seastars to feed locally, reducing their exposition to risks. Moreover, preying without selecting for a specific mussel size did not result in a decrease of net profit, rather it was increased. We concluded that under certain circumstances, not selecting prey results in an increase of net profit at prey size.We discuss our results in the context of the mussel industry where the effect of prey selection on the prey population dynamics is of importance. Mussel size selection by seastars, as described in previous works, will impact the size distribution of mussels within cultured populations affecting growth of the population and reproductive output among others, thus increasing the impact of seastar predation beyond that of just removing mussels from the population. However, this study shows that seastars prey upon availability at high densities, typically encountered in cultivated populations, and therefore their effect on mussel population is restricted to removal of mussels without affecting the size distribution of the population.

A Midsummer Night’s Diet: Snapshot on Trophic Strategy of the Alpine Salamander, Salamandra atra

Information on the trophic ecology of the Alpine salamander, Salamandra atra, is scattered and anecdotal. We studied for the first time the trophic niche and prey availability of a population from an area located in Italian Dolomites during the first half of August. Considering that S. atra is a typical nocturnal species, we collected food availability separately for diurnal and nocturnal hours. Our aims were: (i) to obtain information on the realized trophic niche; (ii) to provide a direct comparison between trophic strategy considering only nocturnal preys or considering all preys; (iii) to study trophic strategy of this species at the individual level. In two samplings nights we obtained prey from 50 individuals using stomach flushing technique. Trophic strategy was determined using the graphical Costello method and selectivity using the relativized electivity index. During the short timeframe of our sample, this salamander showed a generalized trophic strategy. The total trophic availability differed significantly from nocturnal availability. Interindividual diet variation is discussed in the light of the optimal diet theory. Finally, we highlighted that considering or not the activity time of the studied taxon and its preys may lead to a conflicting interpretation of the trophic strategies.

Individual–resource networks reveal distinct fruit preferences of selective individuals from a generalist population of the Helmeted Manakin

Ecological studies traditionally assume that generalist populations are homogeneous in the use of food resources, but empirical evidence supports that intraspecific differences in morphology, physiology and behaviour affect foraging decisions and promote diet variation among individuals. Furthermore, the temporal availability of resources may shape the dynamics of population trophic niche, which ultimately depends on individual niches. In this study, we investigated the seasonal changes in individual‐based networks between the Helmeted Manakin Antilophia galeata, a generalist frugivorous bird, and fruiting plants, following theoretical models of interindividual diet variation based on the Optimal Diet Theory. Selective individuals were the majority of the generalist population of the Helmeted Manakin. Our results suggest that the structure of the individual‐resource networks varied seasonally. We found that modularity was higher than expected by chance in the wet season, when fruit availability was also higher. In the dry season, modules disappeared and the network became more nested. These findings are consistent with the Distinct Preference Model of diet variation. We suggest that downscaling ecological networks to the individual level may reveal emergent properties that, albeit existent, are not evident in species–resources networks.

Interindividual variations in plant and fruit traits affect the structure of a plant-frugivore network

Frugivores select their food in a hierarchical way, from plants to individual fruits, to meet their nutritional requirements. According to the optimal diet theory, finding, handling, and digesting fruits is costly, thus plant species that increase attractiveness and reward are usually preferred by frugivores. The same should be expected for individual plants of the same population, which differ from one another in traits related to frugivore attraction. We tested the hypothesis that plant traits that increase attractiveness and reward to frugivores would be strongly selected by birds in a population of Henriettea succosa (Melastomataceae). In 20 h of focal observation in 19 individual trees (380 h in total), we measured plant and fruit traits known to influence frugivore attraction and reward: plant height, fruit size, and fruit sugar content. In addition, we recorded bird behaviour during fruit consumption. We built two weighted networks of birds and individual plants: one monolayer network and a multilayer network with four layers, one for each type of behaviour. First, we evaluated three weighted descriptors of network structure: nestedness, modularity, and specialization. Then, we calculated metrics of centrality and correlated them with plant traits. We recorded 271 visits by 22 bird species of eight families. The network is modular and specialized, showing that subgroups of H. succosa trees with different trait combinations attract different subsets of bird species, in a way that specialist trees are not connected to a subset of the bird species that visit generalist trees. We also found that centrality metrics reached higher scores in plants with lower height, larger fruits, and intermediate sucrose content. Fruit handling was the predominant foraging behaviour in the multilayer network and represented 90 percent of the interactions. Downscaling a plant-frugivore network to its individual trees showed that the structure of the system is influenced by interindividual variations in the tree population, in which individuals with the best combination of traits occupied central positions in the network.

Network analyses support the role of prey preferences in shaping resource use patterns within five animal populations

Individual variation is an inherent aspect of animal populations and understanding the mechanisms shaping resource use patterns within populations is crucial to comprehend how individuals partition resources. Theory predicts that differences in prey preferences among consumers and/or differences in the likelihood of adding new resources to their diets are key mechanisms underlying intrapopulation variation in resource use. We developed network models based on optimal diet theory that simulate how individuals consume resources under varying scenarios of individual variation in prey preferences and in the willingness of consuming alternate resources. We then investigated how the structure of individual–resource networks generated under each model compared to the structure of observed networks representing five classical examples of individual diet variation. Our results support the notion that, for the studied populations, individual variation in prey preferences is the major factor explaining patterns in individual–resource networks. In contrast, variation in the willingness of adding prey does not seem to play an important role in shaping patterns of resource use. Individual differences in prey preferences in the studied populations may be generated by complex behavioral rules related to cognitive constraints and experience. Our approach provides a pathway for mapping foraging models into network patterns, which may allow determining the possible mechanisms leading to variation in resource use within populations.

A mammoth mouthful? A test of the idea that larger animals ingest larger seeds

AbstractAimIt has been widely assumed that large seeds generally require large animals to ingest and disperse them. However, this relationship has only been quantified in single animal groups (e.g. birds) and in a few communities. Our goal was to provide the first broad‐scale study of the relationship between animal body mass and ingested seed size.LocationGlobal.MethodsWe compiled a dataset of 13,135 unique animal × seed interactions, animal body masses and seed sizes in these interactions, across all vertebrate groups (fish, amphibians, reptiles, birds and mammals).ResultsContrary to expectations, ingested seed size was negatively related to animal body mass. This negative relationship was largely driven by large ungulates ingesting small and dry seeds, and analyses excluding either ungulates or seeds with non‐fleshy fruit types showed a positive relationship between animal body mass and ingested seed size. Large animals ingested both seeds with a larger maximum size (the 95th quantile had a positive slope) and a smaller minimum size (the 5th quantile had a negative slope). Larger animals ingest larger seeds from fleshy fruits but smaller seeds from non‐fleshy fruits. A significant positive relationship was found between animal size and the number of seed species ingested.Main conclusionsOur data show that one of the assumptions that has underpinned the study of animal–seed interactions does not hold true across the full range of animal taxa and fruit types. These findings shed new light on theories about which types of plant species might be at risk if large animals go extinct, and cast doubt on the generality of a few theories (e.g. optimal diet theory, fruit‐size hypothesis) about the relationship between frugivores and seeds.

Individual specialization to non-optimal hosts in a polyphagous marine invertebrate herbivore.

Factors determining the degree of dietary generalism versus specialism are central in ecology. Species that are generalists at the population level may in fact be composed of specialized individuals. The optimal diet theory assumes that individuals choose diets that maximize fitness, and individual specialization may occur if individuals' ability to locate, recognize, and handle different food types differ. We investigate if individuals of the marine herbivorous slug Elysia viridis, which co-occur at different densities on several green macroalgal species in the field, are specialized to different algal hosts. Individual slugs were collected from three original algal host species (Cladophora sericea, Cladophora rupestris and Codium fragile) in the field, and short-term habitat choice and consumption, as well as long-term growth (proxy for fitness), on four algal diet species (the original algal host species and Chaetomorpha melagonium) were studied in laboratory experiments. Nutritional (protein, nitrogen, and carbon content) and morphological (dry weight, and cell/utricle volume) algal traits were also measured to investigate if they correlated with the growth value of the different algal diets. E. viridis individuals tended to choose and consume algal species that were similar to their original algal host. Long-term growth of E. viridis, however, was mostly independent of original algal host, as all individuals reached a larger size on the non-host C. melagonium. E. viridis growth was positively correlated to algal cell/utricle volume but not to any of the other measured algal traits. Because E. viridis feeds by piercing individual algal cells, the results indicate that slugs may receive more cytoplasm, and thus more energy per unit time, on algal species with large cells/utricles. We conclude that E. viridis individuals are specialized on different hosts, but host choice in natural E. viridis populations is not determined by the energetic value of seaweed diets as predicted by the ODT.

The effects of a competitor on the foraging behaviour of the shore crab Carcinus maenas.

Optimal Diet Theory suggests that individuals make foraging decisions that maximise net energy intake. Many studies provide qualitative support for this, but factors such as digestive constraints, learning, predation-risk and competition can influence foraging behaviour and lead to departures from quantitative predictions. We examined the effects of intraspecific competition within a classic model of optimal diet – the common shore crab, Carcinus maenas, feeding on the mussel, Mytilus edulis. Unexpectedly, we found that breaking time (Tb), eating time (Te), and handling time (Th) all decreased significantly in the presence of a conspecific. Reduced handling time in the presence of a competitor resulted in an increased rate of energy intake, raising the question of why crabs do not always feed in such a way. We suggest that the costs of decreased shell breaking time may be increased risk of claw damage and that crabs may be trading-off the potential loss of food to a competitor with the potential to damage their claw whilst breaking the shell more rapidly. It is well documented that prey-size selection by crabs is influenced by both the risk of claw damage and competition. However, our results are the first to demonstrate similar effects on prey handling times. We suggest that crabs maximise their long-term rate of energy intake at a scale far greater than individual foraging events and that in order to minimise claw damage, they typically break shells at a rate below their maximum. In the presence of a competitor, crabs appear to become more risk-prone and handle their food more rapidly, minimising the risk of kleptoparasitism.

The role ofApodemusmice andMicrotusvoles in the diet of the Tengmalm's owl in Central Europe

AbstractBased on a long‐term dataset (1999–2010), we investigated how the availability of main prey affects the breeding density and food ecology of the Tengmalm's owl (Aegolius funereus) in the Czech Republic. In particular, we assessed the role ofMicrotusvoles andApodemusmice in the diet, based on the main predictions of the optimal diet theory that the diet composition depends on the availability of the main prey. We found that (i) the Tengmalm's owl exhibited no numerical response to the availability ofMicrotusvoles andApodemusmice in the field; (ii) the availability ofApodemusmice in the field positively affected their proportion in the diet (26 %), and despite a high proportion ofMicrotusvoles in the owls’ diet (47 %), no relationship was found between their availability in the field and proportion in the diet; (iii) the proportion ofApodemusmice was negatively correlated to the proportion ofMicrotusvoles,Sorexshrews and birds in the diet, but no similar relationship was detected forMicrotusvoles; (iv) the reproductive output of Tengmalm's owls was positively correlated to the proportion ofApodemusmice in the diet, as well as toApodemusmice andMicrotusvole availability in the field; and (v) diet diversity and diet overlap were not significantly affected by the abundance ofApodemusmice andMicrotusvoles. Therefore, the validity of these main optimal diet theory predictions was not confirmed, especially forMicrotusvole prey, due to an opportunistic choice betweenApodemusmice andMicrotusvoles. We suggest that the reproductive output of nocturnal raptors in Central Europe may be less dependent onMicrotusvole supply than that of their northern counterparts.

Diet choice of a predator in the wild: overabundance of prey and missed opportunities along the prey capture sequence

Optimal diet theory (ODT) postulates that predators adjust their foraging decisions by calculating a prey value from the potential biomass gain, handling time, prey vulnerability and encounter rate. Tests of ODT have however so far mainly been restricted to laboratory settings. By video surveillance, we gathered a large data set of more than 2000 field observations of crab spider (Misumena vatia) encounters with potential prey. We then tested whether the complex ODT or two simpler models (prey identity and prey traits) best explain foraging decisions. Insect prey were killed with an average chance of 3.5% when alighting on an inflorescence harboring a spider. Spiders refused to attack suitable prey in 46–79% of encounters when prey was in attack range, indicating an over-abundance of prey relative to the needs of the spiders. Reduction of opportunities to capture prey along the prey capture sequence differed among pollinator groups, with syrphids and solitary bees showing strong avoidance of spiders early in the sequence and bumblebees resisting the final strike. Simple prey traits explained foraging decisions better than ODT, which was not supported. In the absence of food limitation, optimality decisions may be less stringent. The over-abundance of prey indicates that, in contrast to current theory, prey encounter rates are not the most important factor driving predator foraging decisions. Our results are highly coherent with those obtained in earlier field studies on patch leaving strategies and predator-prey encounters. Prey over-abundance and non-optimal predator behavior are apparently not uncommon in nature, and we highlight some of the implications for predator-prey theory.

Diet and energetic constraints of an earthworm specialist, the Mesa Central Blotched Garter Snake (Thamnophis scaliger)

Optimal diet theory predicts that predators optimize energy intake by balancing costs and benefits of foraging. One extreme strategy of snake foraging ecology is shown by specialist species that forage on low-energy prey, such as Thamnophis scaliger (Jan, 1863) which feeds almost exclusively on earthworms. Compared with other prey types such as small mammals, lizards, or arthropods, earthworms are low-energy prey because of their small size and high water content. Given the importance of energy acquisition for fueling snake reproduction, we expect that a low-energy dietary specialist such as T. scaliger needs to forage frequently to store enough fat to reproduce. The high frequency of snakes containing prey, the presence of multiple earthworms in snakes, and the fact that females continue to feed when gravid suggest that T. scaliger is a voracious consumer of earthworms. Despite these foraging behaviours, females did not reproduce in sequential years, suggesting constraints in energy input to reproduce more frequently. A meta-analysis of the diet, body size, and reproductive frequency of some species of the genus Thamnophis Fitzinger, 1843 confirms that consumption of invertebrate prey is associated with small snake size, but not with biennial reproductive frequency within the genus.

The nested assembly of individual-resource networks

1. Much of the current understanding of ecological systems is based on theory that does not explicitly take into account individual variation within natural populations. However, individuals may show substantial variation in resource use. This variation in turn may be translated into topological properties of networks that depict interactions among individuals and the food resources they consume (individual-resource networks). 2. Different models derived from optimal diet theory (ODT) predict highly distinct patterns of trophic interactions at the individual level that should translate into distinct network topologies. As a consequence, individual-resource networks can be useful tools in revealing the incidence of different patterns of resource use by individuals and suggesting their mechanistic basis. 3. In the present study, using data from several dietary studies, we assembled individual-resource networks of 10 vertebrate species, previously reported to show interindividual diet variation, and used a network-based approach to investigate their structure. 4. We found significant nestedness, but no modularity, in all empirical networks, indicating that (i) these populations are composed of both opportunistic and selective individuals and (ii) the diets of the latter are ordered as predictable subsets of the diets of the more opportunistic individuals. 5. Nested patterns are a common feature of species networks, and our results extend its generality to trophic interactions at the individual level. This pattern is consistent with a recently proposed ODT model, in which individuals show similar rank preferences but differ in their acceptance rate for alternative resources. Our findings therefore suggest a common mechanism underlying interindividual variation in resource use in disparate taxa.

Large body and small brain and group sizes are associated with predator preferences for mammalian prey

Predation is a major force in shaping biological communities, both over ecological and evolutionary timescales. In response to predation pressure, prey have evolved characteristics designed to mitigate predation pressure. We evaluated predator foraging biases in relation to prey characteristics across 16 vertebrate communities. We show that although predator biases vary, some prey traits are consistently associated with predator diet composition. Within their acceptable prey size range, predators show positive bias toward larger bodied prey, small-brained prey (controlling for body size), small group size, and terrestriality. Thus, whether predator foraging decisions are passive or active, predator choice exerts differential pressure on prey species according to prey characteristics. Predator biases also were positively associated with early age at maturity, supporting the role of mortality in driving life-history characteristics. These results support several theoretical models of predation including its role as a selective force driving evolutionary changes in life history, brain size and sociality, optimal diet theory, and antiapostatic predation.

Effects of reinforcer delay and variability in a successive-encounters procedure.

Pigeons responded in a successive-encounters procedure that consisted of a search period, a choice period, and a handling period. The search period was either a fixed-interval or a mixed-interval schedule presented on the center key of a three-key chamber. Upon completion of the search period, the center key was turned off and the two side keys were lit. A pigeon could either accept a delay followed by food (by pecking the right key) or reject this option and return to the search period (by pecking the left key). During the choice period, a red right key represented the long alternative (a long handling delay followed by food), and a green right key represented the short alternative (a short handling delay followed by food). The experiment consisted of a series of comparisons for which optimal diet theory predicted no changes in preference for the long alternative (because the overall rates of reinforcement were unchanged), whereas the hyperbolic-decay model predicted changes in preference (because the delays to the next possible reinforcer were varied). In all comparisons, the results supported the predictions of the hyperbolic-decay model, which states that the value of a reinforcer is inversely related to the delay between a choice response and reinforcer delivery.

Testing competing measures of profitability for mobile resources

Optimal diet theory often fails to predict a forager's diet choice when prey are mobile. Because they escape or defend themselves, mobile prey are likely to increase the forager's handling time, thereby decreasing its fitness gain rate. Many animals have been shown to select their prey so as to maximize either their fitness gain or their fitness gain rate. However, no study has yet compared directly these two measures of profitability by generating testable predictions about the choice of the forager. Under laboratory conditions, we compared these two measures of profitability, using the aphid parasitoid Aphidius colemani and its host, Myzus persicae. Fitness gain was calculated for parasitoids developing in each host instar by measuring life-history traits such as developmental time, sex ratio and fecundity. Fitness gain rate was estimated by dividing fitness gain by handling time, the time required to subdue the host. Fourth instar aphids provided the best fitness gain to parasitoids, whereas second instar aphids were the most profitable in terms of fitness gain rate. Host choice tests showed that A. colemani females preferred second instar hosts, suggesting that their decision maximizes fitness gain rate over fitness gain. Our results indicate that fitness gain rate is a reliable predictor of animal's choice for foragers exploiting resources that impose additional time cost due to their mobility.

Discrimination abilities for nutrients: which difference matters for choosy birds and why?

Fine-tuned discrimination abilities are a prerequisite for optimal diet theory (ODT) and the concept of nutrient regulation as alternative models of food choice. These abilities have rarely been determined, and neither has been applied to frugivorous consumers. We investigated nutrient choices of four tanagers (Tachyphonus cristatus, Dacnis cayana, Chlorophanes spiza and Cyanerpes nitidus) for different lipid or carbohydrate concentrations, and determined discrimination abilities by reducing in steps the difference between two foods ranging from 2.5 to 12%. Three species detected differences in sugar concentrations of only 1% and differences in lipid content of 2%. Hence, frugivorous tanagers were able to select the more rewarding food based on fine-scale differences in nutritional content. Tanagers also consumed more carbohydrates than proteins or lipids in isocaloric trials, but were indifferent to equicaloric solutions of 20% of either glucose or sucrose. In contrast to foods with low sugar contents, intake of foods high in lipids or carbohydrates was limited to a protein intake of 0.75g/7h. Lipid and carbohydrates were treated as though they were interchangeable nutrients for birds at low, but not at high, sugar concentrations. Furthermore, the tanagers discriminated foods containing the same amount of protein but differing in the type of protein. Preference for boviserine serum albumin (BSA) over casein was not related to moult. We supplemented the food consisting of casein with the essential amino acid cysteine to match cysteine concentrations of BSA. The birds then consumed more of the casein food than of the cysteine-supplemented casein food. Overall, the birds' consistent choices in the experiments with foods of different energy density are interpretable by both ODT and the nutrient regulation theory. However, only the latter accounts for the birds' choices of isocaloric foods differing in macro- or micronutritional content. Models of food selection should therefore focus on nutrient geometry and consumer-specific requirements. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Effects of the availability of herbaceous food on vole attacks on birch seedlings

According to the optimal diet theory, predators should consume low-quality prey only when the availability of better-quality prey is low. The model of short-term apparent competition predicts, somewhat contradictorily, that low-quality prey co-occurring in the same patches with high-quality prey should be attacked more than low-quality prey in patches containing no high-quality prey. The prediction should apply, in particular, to poor environments. We compared vole attacks on experimental birch seedlings (low-quality prey) in patches with and without herbs (high-quality prey) in experimental enclosures with non-depleted or depleted herbaceous vegetation. Voles attacked fewer birch seedlings in the non-depleted enclosures compared to the depleted ones. They attacked more birch seedlings in patches with herbs than in patches without herbs independently of food depletion in the enclosure. Tree-seedling predation increased with vole density, but the increase was more pronounced in the depleted enclosures. These results may be useful for explaining and controlling natural predation of tree seedlings by voles.