Mate choice decision rules: Trait synergisms and preference shifts

An important and understudied question in sexual selection is how females evaluate information from multiple secondary sexual traits (SSTs), particularly when expression of traits is phenotypically uncorrelated. We performed mate choice experiments on zebra finches (Taeniopygia guttata castanotis Gould) to evaluate two hypotheses: preference shifts (obstacles to choice using one trait increase chooser reliance on others) and trait synergisms (choice based on the sum/product of two or more independently varying traits). The first experiment, which employed males raised on diets that impact SST expression, supported the trait synergism hypothesis: overall, male pairing success was best predicted by synergisms involving beak color and cheek patch size. Results did not support the preference shift hypothesis. Results of a follow‐up experiment that included males reared on a single diet, and in which male beak color and cheek patch size were manipulated, were also consistent with the trait synergism hypothesis. Results have implications for understanding the long‐term persistence of multiple SSTs in populations and for the measurement of repeatability and heritability of mate preferences.     

Phenotypic Correlates of Scrounging Behavior in Zebra Finches: Role of Foraging Efficiency and Dominance

In flocks, individuals can search for their own food using the producer tactic or exploit the discoveries of companions using the scrounger tactic. Models of the producer–scrounger game usually assume that tactic payoffs are independent of individual phenotypic traits. However, factors such as dominance status or foraging efficiency may constrain the use of tactics and lead to asymmetric tactic use among individuals. For instance, in flocks composed of foragers with unequal foraging efficiency, foragers that are less efficient at obtaining food are expected to rely on the scrounger tactic to a greater extent. I examined the role of foraging efficiency and dominance status as potential correlates of scrounging behavior in small aviary flocks of zebra finches (Taenopygia guttata). Individual foraging efficiency was documented in each flock in a treatment that prevented scrounging. In a subsequent treatment that allowed scrounging, higher levels of scrounging occurred as predicted in foragers with lower foraging efficiency. Dominance status was a poor predictor of tactic choice. Birds that arrived later on the foraging grid foraged less efficiently when scrounging was prevented and used scrounging to a greater extent when allowed, suggesting a link between boldness, foraging efficiency and the choice of foraging tactics in small flocks of zebra finches.     

Foraging rate versus sociality in the starling Sturnus vulgaris

It is well established that social conditions often modify foraging behaviour, but the theoretical interpretation of the changes produced is not straightforward. Changes may be due to alterations of the foraging currency (the mathematical expression that behaviour maximizes) and/or of the available resources. An example of the latter is when both solitary and social foragers maximize rates of gain over time, but competition alters the behaviour required to achieve this, as assumed by ideal free distribution models. Here we examine this problem using captive starlings Sturnus vulgaris. Subjects had access to two depleting patches that replenished whenever the alternative patch was visited. The theoretical rate-maximizing policy was the same across all treatments, and consisted of alternating between patches following a pattern that could be predicted using the marginal value theorem (MVT). There were three treatments that differed in the contents of an aviary adjacent to one of the two patches (called the 'social' patch). In the control treatment, the aviary was empty, in the social condition it contained a group of starlings, and in a non-specific stimulus control it contained a group of zebra finches. In the control condition both patches were used equally and behaviour was well predicted by the MVT. In the social condition, starlings foraged more slowly in the social than in the solitary patch. Further, foraging in the solitary patch was faster and in the social patch slower in the social condition than in the control condition. Although these changes are incompatible with overall rate maximization (gain rate decreased by about 24% by self-imposed changes), if the self-generated gain functions were used the MVT was a good predictor of patch exploitation under all conditions. We discuss the complexities of nesting optimal foraging models in more comprehensive theoretical accounts of behaviour integrating functional and mechanistic perspectives.     

Effect of Group Size on Feeding Rate when Patches are Exhaustible

One benefit of group foraging is that individual foragers can join the food discoveries of companions and thus increase encounter rate with food patches. When food patches are exhaustible, however, individual shares of each patch will decrease with group size negating the effect of increased encounter rate. Mean feeding rate may actually decrease with group size as a result of aggression or time wasted joining already depleted patches, or when searching to join the food discoveries of others, which is referred to as scrounging, precludes finding food. I examined the relationship between mean feeding rate and group size in captive flocks of zebra finches (Taenopygia guttata) foraging for small clumps of seeds. Finches in groups of two or four fared better than solitary birds in terms of mean feeding rate despite the fact that birds in groups scrounged a large proportion of their food. Solitary birds initiated feeding activity after a longer delay, which led to their lower success. Early departures by food finders from food patches joined by others may have lessened the impact of scrounging on mean feeding rate. As a result of benefits from the presence of companions, group foraging in zebra finches appears a viable alternative to foraging alone despite the cost of sharing resources.     

Assessing the repeatability,robustness to disturbance,and parent–offspring colony resemblance of collective behavior

Groups of animals possess phenotypes such as collective behaviour, which may determine the fitness of group members. However, the stability and robustness to perturbations of collective phenotypes in natural conditions is not established. Furthermore, whether group phenotypes are transmitted from parent to offspring groups with fidelity is required for understanding how selection on group phenotypes contributes to evolution, but parent–offspring resemblance at the group level is rarely estimated. We evaluated the repeatability, robustness to perturbation and parent–offspring resemblance of collective foraging aggressiveness in colonies of the social spider Anelosimus eximius. Among‐colony differences in foraging aggressiveness were consistent over time but changed if the colony was perturbed through the removal of individuals or via individuals’ removal and subsequent return. Offspring and parent colony behaviour were correlated at the phenotypic level, but only once the offspring colony had settled after being translocated, and the correlation overlapped with zero at the among‐colony level. The parent–offspring resemblance was not driven by a shared elevation but could be due to other environmental factors. The behaviour of offspring colonies in a common garden laboratory setting was not correlated with the behaviour of the parent colony nor with the same colony's behaviour once it was returned to the field. The phenotypes of groups represent a potentially important tier of biological organization, and assessing the stability and heritability of such phenotypes helps us better understand their role in evolution.     

Does Foraging Behaviour Affect Female Mate Preferences and Pair Formation in Captive Zebra Finches?

Background

Successful foraging is essential for survival and reproductive success. In many bird species, foraging is a learned behaviour. To cope with environmental change and survive periods in which regular foods are scarce, the ability to solve novel foraging problems by learning new foraging techniques can be crucial. Although females have been shown to prefer more efficient foragers, the effect of males'' foraging techniques on female mate choice has never been studied. We tested whether females would prefer males showing the same learned foraging technique as they had been exposed to as juveniles, or whether females would prefer males that showed a complementary foraging technique.

Methodology/Principal Findings

We first trained juvenile male and female zebra finches (Taeniopygia guttata) to obtain a significant proportion of their food by one of two foraging techniques. We then tested whether females showed a preference for males with the same or the alternative technique. We found that neither a male''s foraging technique nor his foraging performance affected the time females spent in his proximity in the mate-choice apparatus. We then released flocks of these finches into an aviary to investigate whether assortative pairing would be facilitated by birds taught the same technique exploiting the same habitat. Zebra finches trained as juveniles in a specific foraging technique maintained their foraging specialisation in the aviary as adults. However, pair formation and nest location were random with regard to foraging technique.

Conclusions/Significance

Our findings show that zebra finches can be successfully trained to be foraging specialists. However, the robust negative results of the conditions tested here suggest that learned foraging specializations do not affect mate choice or pair formation in our experimental context.     

A Test of Inbreeding Avoidance in the Zebra Finch

Optimal-outbreeding theory suggests that the genetic relatedness of individuals may be an important factor in mate choice. Several studies have demonstrated that matings between individuals of intermediate genetic relatedness tend to produce more offspring than those between more closely or distantly related mates. Inbreeding avoidance has been studied in captive zebra finches, Poephila guttata. Slater & Clements (1981) found a preference for closely related individuals, while Evans-Layng (1987), in a different design, found a preference for less related mates. Our study examines patterns of mate choice in small flocks of zebra finches of known genetic relatedness. 13 flocks of 9 birds (3 females and 6 males) were released into separate aviaries and left until pairs had formed. 6 of these flocks consisted of siblings and nonrelatives and 7 consisted of cousins and nonrelatives. The degree of genetic relatedness of the individuals used did not affect mate choice, while factors such as male age and experience were correlated with mating success. There was a marginally significant trend for sibling pairs to take longer to lay eggs, although no overall difference in clutch size among sibling pairs, cousin pairs and genetically unrelated pairs was detected. We conclude, then, that there is no evidence of inbreeding avoidance in captive zebra finches.     

Assortative pairing and divergent evolution in Darwin’s Small Tree Finch, Camarhynchus parvulus

Size assortative mating has received increasing attention due to its potential to drive divergence and perhaps speciation. In this study, we examined assortative pairing at 17 nests of Darwin’s Small Tree Finches, Camarhynchus parvulus. We found positive assortative pairing for two traits: bill length and tarsus length, and these traits showed a significant positive correlation to each other. Assortative pairing could be driven by female choice for similar phenotypes because male–male competition has rarely been observed in the Small Tree Finch, nor have males been observed to reject potential mates. Given the high heritability of bill morphology in Darwin’s finches, it is possible that female preference for male bill length, a trait that is known to be important for foraging, will influence offspring bill size to maximise efficient exploitation of resources. The finding of size assortative pairing on the basis of tarsus length requires more research, but suggests different trait utilities for different foraging niches. Interestingly, the highland distribution of tarsus length across the population showed a unimodal distribution, but a bimodal distribution after pairing. While not significant, we found comparatively large differences across study plots in tarsus length, which suggests the possibility of phenotype–habitat matching at a small spatial scale in this species. Our findings are significant in the context of the adaptive radiation of Darwin’s finches as they are consistent with the allopatric model of speciation but also show potential for adaptive divergence in sympatry in Darwin’s tree finches.     

Testing dynamic variance-sensitive foraging using individual differences in basal metabolic rates of zebra finches

Social foragers can alternate between searching for food (producer tactic), and searching for other individuals that have located food in order to join them (scrounger tactic). Both tactics yield equal rewards on average, but the rewards generated by producer are more variable. A dynamic variance-sensitive foraging model predicts that social foragers should increase their use of scrounger with increasing energy requirements and/or decreased food availability early in the foraging period. We tested whether natural variation in minimum energy requirements (basal metabolic rate or BMR) is associated with differences in the use of producer–scrounger foraging tactics in female zebra finches Taeniopygia guttata . As predicted by the dynamic variance-sensitive model, high BMR individuals had significantly greater use of the scrounger tactic compared with low BMR individuals. However, we observed no effect of food availability on tactic use, indicating that female zebra finches were not variance-sensitive foragers under our experimental conditions. This study is the first to report that variation in BMR within a species is associated with differences in foraging behaviour. BMR-related differences in scrounger tactic use are consistent with phenotype-dependent tactic use decisions. We suggest that BMR is correlated with another phenotypic trait which itself influences tactic use decisions.     

The Importance of Validating Experimental Setups: Lessons from Studies of Food Choice Copying in Zebra Finches

The use of social information for foraging decisions has attracted much research attention. One line of research makes use of experimental assays of food choice copying to study whether specific traits of demonstrators influence whether or not an onlooking individual will copy their behavior. For instance, Benskin (Animal Behaviour, 64, 2002 and 823) reported that captive juvenile zebra finches always copied the foraging decision made by a male wearing red color rings as opposed to one wearing green rings. Here, we report an attempt to examine the generalizability of this finding using another zebra finch population. Our experiment did not show any effect of male ring color on onlooker's food choice and further revealed various population‐specific differences in behavior, for instance, in aversion toward novel feeding sources. We therefore conducted two follow‐up experiments to test the validity of the behavioral assay. These experiments revealed that zebra finches of our population often do not copy demonstrator food choice at all, and that copying, if it occurs, may be sensitive to very specific conditions of the experimental setup. Problematically, after testing a total of 124 onlookers (60 in the first experiment, 32 in the second, and 32 in the third), we still cannot confidently say whether or when our birds copy from demonstrators. Hence, we emphasize the risks of adopting experimental procedures that have proven successful in one study, without extensively validating them for the own study. The seemingly more rewarding follow‐up experiments can bear high risks of yielding false‐positive findings that can then be misinterpreted as justification for an actually uninformative experimental procedure.     

Heritability and physiological correlates of migratory tendency in the grasshopper Melanoplus sanguinipes

Abstract. Durations of tethered flights by the North American migratory grasshopper Melanoplus sanguinipes Fabricius are bimodally distributed: most individuals either will not fly, or else will fly for many hours. This observation suggests a simple measure (the ‘one‐hour rule’) for distinguishing migrants from non‐migrants. This measure is repeatable (repeatability = 0.6–0.7). Using laboratory‐reared offspring of grasshoppers from an Arizona population of mixed migratory tendency, a breeding experiment was conducted to determine the heritability of migratory tendency and possible correlated responses to selection on migratory behaviour. When migratory tendency is considered as a threshold trait, the heritability of liability is in the range 0.5–0.6. Most families in the breeding experiment had at least some migrants among their offspring; selection on migratory incidence had a correlated effect on the durations of flights by these individuals. The magnitude of thoracic lipid reserves showed a modest correlated response to selection on migratory behaviour. Thoracic and abdominal lipid reserves in identified migrants are reduced by flight, indicating that lipid is mobilized and consumed during flight in this species.     

Effects of habitat complexity,dominance and personality on habitat selection: Ideal despotic cichlids

Habitat structure can impede visibility and movement, resulting in lower resource monopolization and aggression. Consequently, dominant individuals may prefer open habitats to maximize resource gain, or complex habitats to minimize predation risk. We explored the role of dominance on foraging, aggression and habitat choice using convict cichlids (Amatitlania nigrofasciata) in a two‐patch ideal free distribution experiment. Groups of six fish of four distinct sizes first competed for shrimp in one‐patch trials in both an open and complex habitat; half the groups experienced each habitat type first. Following these one‐patch trials, each group then chose between habitat types in a two‐patch trial while competing for food. Finally, each fish underwent an individual behavioural assessment using a battery of “personality” tests to determine if behaviour when alone accurately reflected behaviour within a social context. In the one‐patch trials, dominant fish showed similar food consumption between habitats, but chased more in the complex habitat. In the two‐patch choice trials, dominants preferred and defended the complex habitat, forming an ideal despotic distribution with more than half the fish and competitive weight in the open habitat. Within the groups, individual fish differed in foraging and chasing, with repeatabilities of 0.45 and 0.23 across all treatments. Although a higher foraging rate during the individual assessment predicted foraging rate and use of the complex habitat during the group trials, aggression and boldness tests were not reflective of group behaviour. Across groups, heavier dominants and those with higher foraging rate in the open habitat used the open habitat more, suggesting that both risk and energetic state affect habitat preference in dominant convict cichlids.     

No evidence of eavesdropping on heterospecific alarm calls by captive zebra finches recently descended from wild birds

Alarm calls are vocalisations animals give in response to predators which mainly function to alert conspecifics of danger. Studies show that numerous species eavesdrop on heterospecific calls to gain information about predator presence. Responding to heterospecific calls may be a learned or innate response, determined by whether the response occurs with or without prior exposure to the call. In this study, we investigated the presence of eavesdropping behaviour in zebra finches Taeniopygia guttata. This species is not known to possess a distinct alarm call to warn adult conspecifics of a threat, and could be relying on alarm calls of nearby heterospecifics for predator information. We used a playback experiment to expose captive zebra finches to three heterospecific sounds: an unfamiliar alarm call (from the chestnut‐rumped thornbill Acanthiza uropygialis), a familiar alarm call, and a familiar control (both from the noisy miner Manorina melanocephala). These calls were chosen to test if the birds had learnt to distinguish between the function of the two familiar calls, and if the acoustic properties of the unfamiliar alarm indicated presence of a threat to the finches. Our results showed that in response to the thornbill alarm, the birds reduced the rate of production of short calls. However, this decrease was also seen when considering both short and distance calls in response to the control sound. An increase in latency to call was also seen after the control stimulus when compared to the miner alarm. The time spent scanning increased in response to all three stimuli, but this did not differ between stimuli. There were no significant differences when considering the stimulus by time interaction for any of the three vigilance measures. Overall, no strong evidence was found to indicate that the captive zebra finches were responding to the heterospecific alarm stimuli with anti‐predator behaviour.     

The influence of differences in social experience on the development of species recognition in zebra finch males

Zebra finch males were first raised by zebra finch parents and then placed in a group of Bengalese finches between the ages of 30 and 60 days. A higher number of aggressive as well as non-aggressive initiatives by Bengalese finches towards young zebra finch males during this period was correlated with a more Bengalese-finch-directed sexual preference when these males were given a choice between a zebra finch and a Bengalese finch female as adults. Experiments in which a zebra finch male was exposed to Bengalese finches behind a wire screen or to Bengalese finch models gave corresponding results. The study shows that, in contrast to earlier findings, zebra finch males raised by their parents for 31 days can still develop a preference for Bengalese finches. Short term changes in preference are discussed. The results indicate that the behaviour shown by stimulus birds in studies on ‘sexual imprinting’ is important for the development of sexual preferences.     

Unrewarding experiences and their effect on foraging in the parasitic waspLeptopilina heterotoma (Hymenoptera: Eucoilidae)

The host-foraging behavior of female entomophagous parasitoids is commonly modified by positive associative learning. Typically, a rewarding experience (e.g., successful oviposition in a host) increases a female's foraging effort in a host microhabitat of the type associated with that experience. Less well understood are the effects of unrewarding experiences (i.e., unsuccessful foraging). The influence of unrewarding experience on microhabitat choice and residence time within a microhabitat was examined for the eucoilid parasitoid,Leptopilina heterotoma, in laboratory and greenhouse assays. As determined previously, females which oviposited successfully in either of two microhabitat types (fermenting apple or decaying mushroom) strongly preferred to forage subsequently on that microhabitat type. However, failure to find hosts in the formerly rewarding microhabitat caused females to reverse their preference in favor of a novel microhabitat type. The effect, though striking, was transient: within 1–2 h, the original learned preference was nearly fully restored. Similar effects of unrewarding experiences were observed with respect to the length of time spent foraging in a microhabitat. As determined previously, oviposition experience in a particular microhabitat type increased the time spent foraging in a patch of that microhabitat type. However, failure to find hosts in the patch caused the time a wasp spent in the next unoccupied patch of that type to decrease to almost nothing. In addition, there was a tendency for an unrewarding experience on a formerly rewarding microhabitat type to extend the time spent in a patch of a novel type. The function of the observed effects of unrewarding experiences is discussed.     

MALE MATE CHOICE AND THE EVOLUTION OF FEMALE PLUMAGE COLORATION IN THE HOUSE FINCH

The house finch (Carpodacus mexicanus) is a sexually dichromatic passerine in which males display colorful plumage and females are generally drab brown. Some females, however, have a subdued version of the same pattern of ornamental coloration seen in males. In previous research, I found that female house finches use male coloration as an important criterion when choosing mates and that the plumage brightness of males is a reliable indicator of male nest attentiveness. Male house finches invest substantially in the care of young and, like females, stand to gain by choosing high-quality mates. I therefore hypothesized that a female's plumage brightness might be correlated with her quality and be the basis for male mate choice. In laboratory mate choice experiments, male house finches showed a significant preference for the most brightly plumaged females presented. Observations of a wild population of house finches, however, suggest that female age is the primary criterion in male choice and that female plumage coloration is a secondary criterion. In addition, yearling females tended to have more brightly colored plumage than older females, and there was no relationship between female plumage coloration and overwinter survival, reproductive success, or condition. These observations fail to support the idea that female plumage coloration is an indicator of individual quality. Male mate choice for brightly plumaged females may have evolved as a correlated response to selection on females to choose brightly colored males.     

Quantitative genetics and behavioural correlates of digit ratio in the zebra finch

A recent study on a captive zebra finch population suggested that variation in digit ratio (i.e. the relative length of the second to the fourth toe) might be an indicator of the action of sex steroids during embryo development, as is widely assumed for human digits. Zebra finch digit ratio was found to vary with offspring sex, laying order of eggs within a clutch, and to predict aspects of female mating behaviour. Hence, it was proposed that the measurement of digit ratio would give insights into how an individual's behaviour is shaped by its maternal environment. Studying 500 individuals of a different zebra finch population I set out to: (1) determine the proximate causes of variation in digit ratio by means of quantitative genetics and (2) to search for phenotypic and genetic correlations between digit ratio, sexual behaviour and aspects of fitness. In contrast to the earlier study, I found no sexual dimorphism in digit ratio and no effect of either laying order or experimentally altered hatching order on digit ratio. Instead, I found that variation in digit ratio was almost entirely additive genetic, with heritability estimates ranging from 71 to 84%. The rearing environment (from egg deposition to independence) explained an additional 5-6% of the variation in digit ratio, but there was no indication of any maternal effects transmitted through the egg. I found highly significant phenotypic correlations (and genetic correlations of similar size) between digit ratio and male song rate (positive correlation) as well as between digit ratio and female hopping activity in a choice chamber (negative correlation). Rather surprisingly, the strength of these correlations differed significantly between subsequent generations of the same population, illustrating how quickly such correlations can appear and disappear probably due to genotype-environment interactions.     

Zebra mussels affect benthic predator foraging success and habitat choice on soft sediments

The introduction of zebra mussels (Dreissena spp.) to North America has resulted in dramatic changes to the complexity of benthic habitats. Changes in habitat complexity may have profound effects on predator-prey interactions in aquatic communities. Increased habitat complexity may affect prey and predator dynamics by reducing encounter rates and foraging success. Zebra mussels form thick contiguous colonies on both hard and soft substrates. While the colonization of substrata by zebra mussels has generally resulted in an increase in both the abundance and diversity of benthic invertebrate communities, it is not well known how these changes affect the foraging efficiencies of predators that prey on benthic invertebrates. We examined the effect of zebra mussels on the foraging success of four benthic predators with diverse prey-detection modalities that commonly forage in soft substrates: slimy sculpin (Cottus cognatus), brown bullhead (Ameirus nebulosus), log perch (Percina caprodes), and crayfish (Orconectes propinquus). We conducted laboratory experiments to assess the impact of zebra mussels on the foraging success of predators using a variety of prey species. We also examined habitat use by each predator over different time periods. Zebra mussel colonization of soft sediments significantly reduced the foraging efficiencies of all predators. However, the effect was dependent upon prey type. All four predators spent more time in zebra mussel habitat than in either gravel or bare sand. The overall effect of zebra mussels on benthic-feeding fishes is likely to involve a trade-off between the advantages of increased density of some prey types balanced against the reduction in foraging success resulting from potential refugia offered in the complex habitat created by zebra mussels.     

Transgenerational effects on body size caused by early developmental stress in zebra finches

The nutritional and social conditions that individuals experience during early development can have profound effects on their morphology, physiology, behaviour and life history. Experimental increases in brood size in birds can result in reduced offspring condition and survival, indicating that developmental deficits in enlarged broods have negative fitness consequences within the affected generation. To study long-term effects (i.e. transgenerational effects of developmental stress), we conducted a two-step breeding experiment in which we manipulated early developmental conditions in zebra finches Taeniopygia guttata. We raised zebra finches by manipulating brood sizes and controlled for maternal and genetic effects by cross-fostering. In a previous study, we showed that offspring condition and body size decreased with increasing brood size. Here we show that this effect was carried over to the next generation. Body size in nestlings and at nutritional independence was affected by the brood size in which the mothers were raised. Female offspring did significantly worse than male offspring when the mother had been raised in large broods, suggesting a sex-specific influence of maternal effects. These findings link early developmental stress in females with the phenotype of the next generation via maternal effects.     

Optimal foraging: food patch depletion by ruddy ducks

Summary I studied the foraging behavior of ruddy ducks (Oxyura jamaicensis) feeding on patchily distributed prey in a large (5-m long, 2-m wide, and up to 2-m deep) aquarium. The substrate consisted of a 4x4 array of wooden trays (1.0-m long, 0.5-m wide, and 0.1-m deep) which contained 6 cm of sand. Any tray could be removed from the aquarium and loaded with a known number of prey. One bird foraged in the aquarium at a time; thus, by removing a food tray after a trial ended and counting the remaining prey, I calculated the number of prey consumed by the bird. I designed several experiments to determine if ruddy ducks abandoned a food patch in a manner consistent with the predictions of a simple, deterministic, patch depletion model. This model is based on the premise that a predator should maximize its rate of net energy intake while foraging. To accomplish this, a predator should only remain in a food patch as long as its rate of energy intake from that patch exceeds the average rate of intake from the environment. In the majority of comparisons, the number of food items consumed by the ruddy ducks in these experiments was consistent with the predictions of the foraging model. When the birds did not forage as predicted by the model, they stayed in the patch longer and consumed more prey than predicted by the model. An examination of the relation between rate of net energy intake and time spent foraging in the food patch indicated that by staying in a patch longer than predicted, the ruddy ducks experienced only a small deviation from maximum rate of net energy intake. These results provided quantitative support for the prediction that ruddy ducks maximize their rate of net energy intake while foraging.