The Effects of Early Feeding Experience on Long-term Seed Choice by Canaries (Serinus canaria)

Early experience has been found to have long-term effects on behaviour such as social imprinting and song learning. In this paper we investigate whether early experience with particular types of food can lead to long-term feeding preferences in canaries. For 8 wk following the onset of independent foraging, two groups of naive fledgling canaries were reared on different diets. One group had a choice of four seed types, while individuals in the other group received only one type of seed. After this training period, all the birds were given a mixed seed diet for a further 15 wk, and then their preference for the original seeds was tested. All six birds that had been given four seeds during the training period showed the same preference, namely: hemp > niger > millet > linseed. However, all the birds that had been trained on one seed type only, still preferred the seed on which they had been trained. After a further 23 wk on a mixed-seed diet, birds from the restricted early diet group showed no further preference for the seed on which they were trained, and their seed choice at this stage was similar to the birds that had received four seeds. We conclude that early experience can affect food choice in canaries for at least 15 wk, but is subordinate to other influences in the longer term.     

Carbon and nitrogen isotopes trace nutrient exchange in an ant-plant mutualism

By combining and reanalysing data from two independent field experiments we explore whether food limitation at the brood stage affects habitat selection in nesting mallards (Anas platyrhynchos). In an introduction experiment we found that, independent of treatment, some study lakes remained empty of wild mallard pairs (”empty lakes”), whereas on other lakes introduced birds attracted wild mallards (”attractive lakes”). In the other experiment we used mallard ducklings to address brood-stage food limitation by studying mass change of ducklings. We found that ducklings foraging on lakes that did not attract wild mallard pairs in the introduction experiment gained much less mass than those foraging on attractive lakes. In most cases ducklings even lost mass in the empty-lake foraging trials, providing strong evidence for food limitation. Therefore, lakes that remained empty of wild mallard pairs in the introduction experiment proved to be inferior brood habitats, particularly in terms of food. Our results give insight into the mechanisms underlying the general habitat selection hypotheses, specifically the ideal preemptive and conspecific attraction rules. The results further support our earlier conclusion that mallards do not use the ideal preemptive rule when selecting nesting lakes. However, conspecific attraction may not be generally applicable either, because, independent of the presence of introduced conspecifics, wild mallards somehow anticipated the low quality of the empty lakes as brood-rearing habitats and made their habitat-selection decision accordingly. Received: 22 March 1999 / Accepted: 7 October 1999     

Mallards Feed Longer to Maintain Intake Rate under Competition on a Natural Food Distribution

Animals foraging in groups may benefit from a faster detection of food and predators, but competition by conspecifics may reduce intake rate. Competition may also alter the foraging behaviour of individuals, which can be influenced by dominance status and the way food is distributed over the environment. Many studies measuring the effects of competition and dominance status have been conducted on a uniform or highly clumped food distribution, while in reality prey distributions are often in‐between these two extremes. The few studies that used a more natural food distribution only detected subtle effects of interference and dominance. We therefore conducted an experiment on a natural food distribution with focal mallards Anas platyrhynchos foraging alone and in a group of three, having a dominant, intermediate or subordinate dominance status. In this way, the foraging behaviour of the same individual in different treatments could be compared, and the effect of dominance was tested independently of individual identity. The experiment was balanced using a 4 × 4 Latin square design, with four focal and six non‐focal birds. Individuals in a group achieved a similar intake rate (i.e. number of consumed seeds divided by trial length) as when foraging alone, because of an increase in the proportion of time feeding (albeit not significant for subordinate birds). Patch residence time and the number of different patches visited did not differ when birds were foraging alone or in a group. Besides some agonistic interactions, no differences in foraging behaviour between dominant, intermediate and subordinate birds were measured in group trials. Possibly group‐foraging birds increased their feeding time because there was less need for vigilance or because they increased foraging intensity to compensate for competition. This study underlines that a higher competitor density does not necessarily lead to a lower intake rate, irrespective of dominance status.     

Taxonomic status of the Forest Buzzard Buteo oreophilus trizonatus

The foraging efficiencies of four sympatric southern African seed-eating birds, namely Bronze Mannikin Spermestes cucullatus, Cape Sparrow Passer melanurus, Southern Red Bishop Euplectes orix and Thick-billed Weaver Amblyospiza albifrons, and a domesticated species, the Bengalese Finch Lonchura domestica, were measured and compared using giving-up densities (the amount of food remaining following patch exploitation) in experimental food patches. Foraging efficiency was quantified using giving-up densities by offering individual birds equal foraging opportunities. A low giving-up density displays the ability of a forager to profitably harvest food at low resource densities and to gainfully exploit the foraging opportunities overlooked by a less efficient forager. Ten individuals of each of the five species were allowed to forage on six different seed types. Thick-billed Weavers had significantly lower giving-up densities for all seed types except the smallest, namely red manna. Bronze Mannikins showed the converse trend, foraging most efficiently on the smallest seeds. The results of the present study revealed that Thick-billed Weavers were the most efficient foragers (i.e. had the lowest giving-up densities on seeds in feeding trays).     

Do Insectivorous Birds use Volatile Organic Compounds from Plants as Olfactory Foraging Cues? Three Experimental Tests

Some insectivorous birds orient towards insect‐defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect‐defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field‐based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore‐damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore‐damaged trees.     

Downy woodpecker foraging behavior: foraging by expectation and energy intake rate

Summary I describe an artificial patch system that was used to study the foraging behavior of free-roaming downy woodpeckers (Picoides pubescens) in a woodlot in southeastern Michigan. The artificial patches used were thin logs into which were drilled small holes to hold food items (bits of sunflower seed kernels). Downy woodpeckers would systematically search the holes of a patch for food items and thus by manipulating the food distribution within the patches, the birds could be made to experience differing rates of energy intake while foraging.Simple deterministic theories of optimal foraging in patchy environments indicate that an optimal forager, who experiences a decreasing rate of energy intake while foraging in a patch, should leave a patch when its rate of energy intake falls below the average intake rate for the overall environment. In other words, an optimal forager is continually assessing the quality of a patch and makes decisions as to when to leave a patch via its energy intake rate. When the downy woodpeckers studied could encounter any one of several types of patches each with differing, decreasing rates of energy intake, they followed a patch quality assessment strategy similar to that suggested by theory. Upon encountering a single type of patch for a number of consecutive days, however, the birds appeared to forage according to prior expectations of patch quality and not according to a quality assessment strategy based on energy intake rates. The observed expectations were not related to the number of food items per patch but they appeared to be based on expectations of when or where to leave a patch.     

Evidence of associative blocking during foraging in the social wasp Vespula germanica

While foraging, Vespula germanica usually return to abundant food sites. During this relocation behavior, these wasps learn to identify contextual cues associated with food position. We analyzed associative blocking in this species, that is, how an association with a conditioned stimulus (CS1) blocks subsequent learning when a novel stimulus (CS2) is added on a second foraging visit. Three groups of wasps (A, B, and C; total 74 individual wasps) were observed while collecting meat during one or two consecutive visits. In group A, an environmental cue (CS1) was paired with food placed at a specific site, and on the second visit, a second cue (CS2) was added while food remained in the same position. In a subsequent testing phase, CS1 was removed and the food source displaced nearby. We then recorded the number of hovers performed over the empty dish (previously baited). Group A wasps appeared to ignore the addition of CS2 on their second visit because they performed fewer hovers over the learned site. For group A, the duration of the decision-making process to finally fly toward the baited dish was shorter than when CS1 and CS2 were presented together on their first visit (group B). This is the first study to demonstrate the occurrence of associative blocking in vespids, confirming that a prior foraging experience influences subsequent food relocation in V. germanica. Our findings reveal that first learning episodes block further associations with novel contextual cues, contributing to understanding of complex cognitive processes involved in V. germanica´s foraging behavior.     

Habitat use by seed-eating birds: a scale-dependent approach

The seedbank in arable farmland represents an important foraging resource for birds, particularly in grassland landscapes where alternative foraging opportunities may be scarce. We used a stepwise approach to examine the importance of seed food resources for farmland birds in winter. First, results from a large-scale experiment in which seed resources were manipulated to test the notion that birds aggregate at food resources, subject to a minimum threshold level. Secondly, a multiscale approach was used to characterize habitat use at a landscape scale and how this may inform agri-environment implementation. Overall seed resources declined sharply over the winter and were relatively low in most arable fields. Large-scale declines in the arable seedbank mean that much habitat may not be of sufficient quality to support foraging bird populations through a winter. At a landscape scale, extensive analyses of breeding season abundance show that bird abundance is most influenced by arable (i.e. seed-rich) habitat in grassland landscapes. The scale at which birds respond to habitat differs between species, and is related to the extent of between-season movements. Implementation of agri-environment schemes will need to consider both the quality of habitat and the context in which it is provided if recent declines in farmland bird populations are to be reversed.     

Leaving final-cut grass silage in situ overwinter as a seed resource for declining farmland birds

The loss of seed-rich wintering habitats has been a major contributory cause of farmland bird population declines in western Europe. Agricultural grasslands are particularly poor winter foraging habitats for granivorous birds, which have declined most in the pastoral farming regions of western Britain. We describe an experiment to test the utility of fertile ryegrass (Lolium) swards as a potentially rich source of winter seed for declining farmland birds. Four patches of final-cut grass silage were allowed to set seed and were left in situ overwinter. Half of each patch was lightly aftermath grazed in an attempt to increase the accessibility of the seed to foraging birds and reduce the perceived predation risk. Large numbers of yellowhammers (Emberiza citrinella) and reed buntings (E. schoeniclus) foraged on the seeded plots throughout the winter. They preferred to forage on ungrazed seeded plots, where the accumulation of senescent foliage resulted in a 14% average loss in silage yield in the following season. However, seed produced on the plots also led to sward regeneration, increasing subsequent yields on some plots. The technique offers clear benefits as a potential future agri-environment measure for declining granivorous birds, with wide applicability, but requires further development to minimise sward damage and costs to the farmer. Autumn grazing should reduce sward damage, but at the cost of reduced usage by buntings. Using the technique just prior to reseeding would be one way of avoiding any costs of sward damage.     

Seed food preferences of granivorous farmland passerines

Capsule Large buntings prefer cereal grains whilst sparrows also take oily seeds.

Aims To determine seed food preferences of Tree Sparrow Passer montanus and Corn Bunting Emberiza calandra.

Methods Forty seed choice experiments were conducted at two sites over two winters. In each experiment, two seed types were provided and the number of visits made by birds to each type was recorded over a set period. At one site, Tree Sparrows were colour-ringed, allowing choices made by individual birds to be recorded. Data were also collected for House Sparrow Passer domesticus, Yellowhammer Emberiza citrinella and Reed Bunting E. schoeniclus.

Results All five species fed intensively on cereal grain, and wheat and oats were consistently preferred to barley. Sparrows and Reed Buntings also took maize, which was avoided by Corn Bunting and Yellowhammer. Tree Sparrow exhibited a broad diet, selecting cereal grain and oily seeds including sunflower and oilseed rape, but rye-grass seed was almost completely avoided.

Conclusion Cereal grain should be a key component of over-winter provision of seed for farmland passerines, especially when targeted at Corn Bunting and Yellowhammer. Oily seeds such as brassicas and sunflower will benefit species with more generalist diets, including Tree Sparrows.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Time budgets of Nuthatches Sitta europaea with supplementary food

Individual time budgets during winter and spring were studied in a population of Nuthatches Sitta europaea in south-central Sweden. In late winter birds spent about 90% of daytime foraging, males feeding somewhat less than females, possibly due to male dominance. In early spring, foraging time decreased to about 65% of daytime activity, and sex-related differences became more pronounced. Males then increased time spent resting and singing. Females were nestbuilding, spent less time resting and also devoted more time to foraging than did males. Prior to the observation periods, food density for part of the population was experimentally increased by supplying sunflower seed. In spite of this, time budgets in the experimental group were similar to those of the control group. It is possible that the intensity of foraging, rather than actual time devoted to it, was influenced by food density and that, as a result, birds in the experimental group were able to maintain a higher level of vigilance. Also, an increase in food supply may be less important to a hoarding bird such as the Nuthatch. However, the result remains that Nuthatch time budgets in winter and spring are not readily influenced by an increase in food density.     

Foraging by little auks in the distant marginal sea ice zone during the chick-rearing period

During the chick-rearing period, little auks Alle alle adopt a bimodal foraging strategy, alternating long trips with several short ones. It has been postulated that they reach more remote areas during long feeding trips than during short ones. However, the range of their foraging flights has never actually been measured. The aims of this study were to find the exact location of the little auk feeding grounds and to investigate whether they reach remote areas during long foraging trips using miniature GPS and temperature loggers. The study was conducted in 2009 in Magdalenefjorden (79°34′N, 11°04′E), one of the main breeding grounds of little auks on Spitsbergen. The temperature logger records indicated that during short trips, little auks visit warmer waters (situated close to the colony) than during long ones. The tracks of two GPS-equipped birds indicated that during long trips little auks foraged in the distant, food-abundant marginal sea ice zone, at least 100 km away from the colony. During long trips, birds make several stops at sea, perhaps sampling the foraging area with respect to prey distribution. Since food conditions near the studied colony are usually suboptimal, little auks may be exploiting distant feeding areas to compensate for the poorer-quality food available at nearby foraging grounds. The extended duration of long foraging trips may enable birds to collect food for chicks on food-abundant, remote foraging grounds as well as acquire, process and excrete food needed for self-maintenance, reducing the costs of flight to the colony.     

White Storks, <Emphasis Type="Italic">Ciconia ciconia</Emphasis>, forage on rubbish dumps in Poland—a novel behaviour in population

Information on the foraging of White Storks on rubbish dumps, a novel behaviour in Central European populations, is presented. Observations were first made in 1999 and to date; they have been recorded on sixteen locations in Poland. From one to 348 White Storks (median = 2, N = 116) were observed on rubbish dumps, and most of the records (86%) were of 1–3 birds. Birds foraging on rubbish dumps were recorded from late March to early September, but the highest numbers were seen during the summer months. Most of the birds (77%, N = 171) were recorded foraging directly on an area where rubbish had been thrown, while a minor fraction (16%) was seen on neighbouring recultivated grassy areas. Dumps were used as an additional food source probably in areas where natural foraging grounds are limited. The increased frequency of observations during the summer months probably resulted from the greater nutritional needs of large nestlings which force the adults to find alternative food sources or foraging habitat shift by non-breeding birds.     

Habitat selection at multiple spatial scales by foraging Glossy Black‐cockatoos

Abstract: Habitat selection among vertebrates entails decision making at a number of spatial scales. An understanding of factors influencing decisions at each of these scales is required for the effective management of wildlife populations. This study investigates the foraging ecology of a population of Glossy Black‐cockatoos in central New South Wales. We took advantage of the characteristic feeding sign produced by Glossy Black‐cockatoos to examine factors influencing habitat selection at multiple spatial scales. Birds preferred to forage at sites where food was abundant and avoided open sites where the predation risk may be greater. Their two food species, Allocasuarina diminuta and Allocasuarina gymnanthera, differed in profitability (kernel intake rate as measured by the ratio of seed weight to total seed and cone weight), as did trees within a species. Both species were utilized extensively, although foraging intensity was greater at sites where the more profitable species was present. In order to maximize their food intake, birds selected individual trees on the basis of cone abundance and profitability. Cones produced in the previous year were preferred.     

African grey parrots (Psittacus erithacus) use inference by exclusion to find hidden food

Exclusion allows the detection of hidden food when confronted with the choice between an empty and a potentially baited food location. However, exclusion may be based on avoidance of the empty location without drawing inferences about the presence of the food in the baited location. So far, such inferences have been demonstrated in the great apes only: after seeing an experimenter eating one of two food types, which both had been hidden previously in two boxes, the apes were able to choose the box that still contained the other food type. African grey parrots are capable of exclusion, and we here assessed if they are capable of inference by exclusion. In our task, two different but equally preferred food items were hidden in full view of the birds under two opaque cups. Then, an experimenter secretly removed one food type and showed it to the bird. Similarly to the apes, one out of seven parrots significantly preferred the baited cup; control conditions rule out that its choice was based on associative learning or the use of olfactory cues. Thus, we conclude that-like the apes-some grey parrots are able to infer the location of a hidden food reward.     

The significance of seed food in chick development re‐evaluated by tracking day‐to‐day dietary variation in the nestlings of a granivorous passerine

The dietary adjustment of nestlings of granivorous birds to a seed diet and the different morphological characteristics of ingested food have rarely been examined in natural conditions. It has been suggested that the provision of cereal grains to nestlings of some seed‐eating bird species in modern agroecosytems is the result of poor food conditions after agricultural intensification. We analysed the abundance of invertebrate prey in the main foraging habitat of parent birds, daily changes (from hatching to fledging) in the efficiency of cereal seed digestion, and the dietary characteristics, diet composition and prey type delivered to nestlings of the Yellowhammer Emberiza citrinella. Analysis of faecal sacs from nests located in breeding habitat with an abundant invertebrate fauna revealed no relationship between the proportion of cereal seeds in the diet of nestlings and the food supply in the main foraging sites of the parents. Neonate nestlings (1 day old) exclusively received weakly chitinized invertebrate prey (arachnids and flies), but from the second day of life the nestlings were fed a variety of highly chitinized invertebrate prey, the percentage biomass of which did not change for the remainder of the nesting period. Cereal grains started to be delivered to 3‐day‐old nestlings and were already efficiently digested, and the percentage biomass of this food type increased progressively with nestling age. We suggest that the provisioning of cereal grains to nestlings is not forced by external factors, such as modern agricultural intensification; rather, it is an intentional behaviour of parent birds aimed at achieving physiological adjustment to seed food in the early stages of ontogeny.     

The effect of unripe fruits on ripe fruit removal by birds in Pistacia terebinthus: flag or handicap?

The shrub Pistacia terebinthus produces crowded infructescences with up to several hundred fruits, which are bright red when unripe and turn green when ripe. Most fruits contain an empty seed and never reach maturity. More ripe fruits were removed by birds from experimental bicolored fruit displays (consisting of infructescences with ten ripe fruits and stripped of unripe fruits, paired with infructescences with only unripe fruits) than from monocolored ones (single infructescences with ten ripe fruits and stripped of unripe fruits). Thus, the presence of unripe fruits seems to increase the conspicuousness or attractiveness of fruit displays to fruit-eating birds. A second experiment compared ripe fruit removal from experimental infructescences having only ripe fruits, with that from control infructescences containing both ripe fruits and natural numbers of unripe fruits, all on P. terebinthus plants. Unlike the first experiment, each bicolored display in this case consisted of a single infructescence with both unripe and ripe fruits. A higher proportion of ripe fruits was removed by birds from infructescences free of unripe fruits. This result suggests that the presence of unripe fruits reduces the accessibility of ripe fruits for fruit-eating birds. This is further supported by field observations of bird foraging behavior.     

Aimed Throwing as a Means of Food Transfer Between Tufted Capuchins (Cebus apella)

We examined aimed throwing as a means of food transfer in tufted capuchins (Cebus apella). We conducted this research in three phases. In Phase 1 we provided food to monkeys in one of two groups housed 1 m apart. We did not provide food to subjects in the second group. An observer recorded each instance in which a subject in the first group threw food toward one in the second group. In Phase 2 we provided a group of capuchins with food and noted each instance in which a subject threw food toward an empty cage. In Phase 3 we provided food simultaneously to two groups of capuchins and noted each instance of food-throwing between them. In Phase 1 subjects in one group threw food toward subjects in a second group, which, when provided the opportunity, did not throw food toward capuchins in the first group. Thrown food was either caught, retrieved, or lost on the test room floor. The rate of throwing decreased significantly when subjects were presented with an empty cage and when both groups of subjects were given food. We propose that psychological processes which underlie aimed throwing and food sharing came into existence through convergent evolution in large-brained, extractive foraging primates. We further speculate that although a well-developed system of exchange, based on contingent reciprocity, may occur among primates only inHomo,simpler transfer systems involving voluntary unidirectional passing of food from one individual to another appear to be more widespread among primates than previously thought and can be expressed in rather unusual circumstances such as those in this experiment.     

Gaze direction - a cue for hidden food in rooks (Corvus frugilegus)?

Other individual's head- and eye-directions can be used as social cues indicating the presence of important events. Among birds, ravens and rooks have been shown to co-orient with conspecifics and with humans by following their gaze direction into distant space and behind visual screens. Both species use screens to cache food in private; also, it had been suggested that they may rely on gaze cues to detect hidden food. However, in an object-choice task, ravens failed to do so, and their competitive lifestyle may have prevented them from relying on these cues. Here we tested closely related and cooperative rooks. Food was hidden in one of two cups and the experimenter gazed at the baited cup. In a second experiment, we aimed to increase the birds’ motivation to choose correctly by increasing the investment needed to obtain the reward. To do so, the birds had to pull on a string to obtain the cup. Here, the birds as a group tended to rely on gaze cues. In addition, individual birds quickly learned to use the cue in both experiments. Although rooks may not use gaze cues to find hidden food spontaneously, they may quickly learn to do so.