Seed selection by dark-eyed juncos (Junco hyemalis): optimal foraging with nutrient constraints?

Summary Observations of the foraging behavior of six captive dark-eyed juncos (Junco hyemalis) are used to test the assumptions and predictions of optimal diet choice models (Pyke et al. 1977) that include nutrients (Pulliam 1975). The birds sequentially encountered single seeds of niger thistle (Guizotia abyssinica) and of canary grass (Phalaris canariensis) on an artificial substrate in the laboratory. Niger thistle seeds were preferred by all birds although their profitability in terms of energy intake (J/s) was less than the profitability of canary grass seeds. Of four nutritional components used to calculate profitabilities (mg/s) lipid content was the only characteristic that could explain the junco's seed preference. As predicted by optimal diet theory the probability of consuming niger thistle seeds was independent of seed abundance. However, the consumption of 71–84% rather than 100% of the seeds encountered is not consistent with the prediction of all-or-nothing selection. Canary grass seeds were consumed at a constant rate (no./s) independent of the number of seeds encountered. This consumption pattern invalidates a model that assumes strict maximization. However, it is consistent with the assumption that canary grass seeds contain a nutrient which is required in minimum amounts to meet physiological demands (Pulliam 1975). These experiments emphasize the importance of incorporating nutrients into optimal foraging models and of combining seed preference studies with studies of the metabolic requirements of consumers.     

Diets of Kauai’s invasive rose-ringed parakeet (<Emphasis Type="Italic">Psittacula krameri</Emphasis>): evidence of seed predation and dispersal in a human-altered landscape

Rose-ringed parakeets (Psittacula krameri) are the world’s most successful introduced parrots, and > 2000 individuals reside on Kauai, Hawaii. These birds destroy crops, but impacts to other native and non-native species are largely unknown. Our study objectives on Kauai were to determine: (1) diets of rose-ringed parakeets at five sites (n = 9–25 per site), by sex, through crop and gizzard analysis and carbon and nitrogen stable isotope analysis, and (2) whether birds are dispersing or depredating seeds. We found 100% of birds (n = 64) were eating plant material and 80% of their diet was seed; males had more food in their crops and gizzards than did females. Corn (Zea mays) was eaten by 67% of birds and averaged 31% of mass in crops and gizzards. Invasive yellow guava (Psidium guajava) was eaten by 97% of birds and averaged 30% of their diet. Parakeets are potentially dispersing yellow guava seeds, as 66% of birds had intact guava seeds, and each bird had an average of three intact seeds. Diets differed statistically among sites. Parakeets from Lihue Airport did not have any corn, and isotopic carbon values also supported low feeding on corn by birds at Lihue Airport. No seeds of native plants were identified in rose-ringed parakeet diets. Our findings of a diverse plant diet, frequent seed predation, and potential to disperse invasive species’ seeds implies that land managers in agricultural, urban, and natural areas should be concerned with the current expansion of these invasive birds on Kauai and elsewhere.     

Avian seed preference and weight loss experiments: the effect of fungal endophyte-infected tall fescue seeds

Summary The impact of endophytic fungus-infected seeds on seed predators is poorly understood. In this multiple trophic level investigation, seed preference experiments were conducted to determine whether five species of passerines (dark-eyed juncos, Junco hyemalis; American tree sparrows, Spizella arborea; song sparrows, Melospiza melodia; chipping sparrows, Spizella pusilla; and house sparrows, Passer domesticus) recognize and preferentially consume noninfected (NI) over infected (I) seeds of tall fescue (Festuca arundinacea). We predicted that the birds would refrain from eating I seeds because those seeds contain high concentrations of fungal alkaloids. When given a choice of NI fescue seeds and control seeds (millet), all bird species showed a significant preference for millet. However, individuals of all species consumed some NI seeds. When given a choice of NI and I fescue seeds, all species except the chippig sparrow ate significatly more NI than I fescue seed and the chipping sparrow showed the same trend. Thus, birds were able to distinguish between the two seed types and preferred NI seeds in choice tests. Additional experiments investigated weight changes in dark-eyed juncos fed diets containing different proportions of millet, NI, and I fescue seed. Significant differences in weight loss were observed for the various diets. Juncos showed greater weight loss when the proportion of fescue seed, especially the proportion of I seed, in their diet was greater. The potential significance of the finding that abundant grass seeds are made unavailable to predators by fungal infection is discussed in relation to foraging and competition in avian communities.     

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 influence of seed packaging and fruit color on feeding preferences of American robins

We investigated the role of seed packaging (division of total seed volume among individual seeds) and fruit color in determining feeding preferences of American Robins (Turdus migratorius). Experiments were conducted using artificial fruits with either 8 small plastic beads or a single large one with equivalent volume. Other fruit characters were held constant. As predicted, large seeds were voided rapidly by regurgitation, resulting in higher pulp consumption rates for large-seeded fruits than for small-seeded ones, whose seeds were passed through the gut. Most birds apparently used this difference in profitability as a choice criterion: four of seven preferred large-seeded fruits. That three individuals did not do so suggests that birds may differ in their ability to perceive minor differences in fruit profitability, or to use them as choice criteria. Pulp color was also important: blue fruits were preferred by all seven birds. This preference was surprising, since Robins commonly feed on red fruits in the field.     

Does Seed‐Caching Experience Affect Spatial Memory Performance by Pinyon Jays?

Food‐storing birds use spatial memory to find previously cached food items. Throughout winter, pinyon jays (Gymnorhinus cyanocephalus) rely heavily on cached pinyon pine (Pinus edulis) seeds. Because of a recent severe drought, pinyon pine trees had not produced a significant seed crop for several years. Therefore, 1‐ and 2‐yr‐old birds never had the opportunity to cache and recover seeds and birds 4 or more years of age had not recovered seeds in 3 yr. This study examined whether natural but extreme variability in experience might result in differences in abstract spatial memory ability during a non‐cache recovery test of spatial memory. Three groups of jays were tested for spatial memory ability in an open room analog of the radial arm maze. Two of the groups were 8 mo old: young/minimally experienced birds which had 2 mo of experience in the wild, while young/experienced birds had 5 mo of experience in their natural habitat. The third group, adult, consisted of birds more than 3 yr old, with at least 3 yr of experience in their natural habitat. This was the only group with experience caching pine seeds. All three groups performed equally and well above chance. This suggests that spatial memory is fully developed by 8 mo of age and is not affected by extensive experience in the wild.     

Regulation of cache stores and body mass in Carolina chickadees (Panes carolinensis)

Here I report the results of laboratory experiments on cachingbehavior of Carolina chickadees (Parus carolinensis) designedto test the following predictions from a recent dynamic optimizationmodel: under limited resources, small birds, when fat, shouldcache food instead of eating it, but they should eat insteadof caching when lean; when resources are abundant, birds shouldcache less when fat and more when lean. In addition, when resourcesare abundant the amount of time spent in foraging-related activityshould decrease with an increase in body mass. Chickadees weretested for 2 weeks in either a poor-quality environment (wherethey were given four 5-min periods per day of access to a feederfilled with sunflower seeds) or a richer environment (wherethey were given four 10-min access periods); they were thenswitched to the alternative environment for an additional 2weeks. The entire experiment lasted from October through June.Within-individual comparisons showed that birds in both thepoor and rich environments exhibited the predicted correlationsbetween the probability that a seed was cached and body massmeasured at dawn. The number of seeds already stored had a weakand variable effect on caching decisions. Time budgets changedas predicted; birds spent less time with food-related behaviorswhen their energy stores were high compared to when they werelow, and overall spent a higher fraction of their time foragingin the poor-quality environment compared to the rich environment.Two other variables also affected caching behavior: experienceand season. Individuals were more likely to cache in the poorenvironment when they had been tested in the rich environmentthe previous 2 weeks, as compared with birds started in thepoor environment. However, this was true only for tests donein the winter. In late spring, all birds stopped caching orreduced caching rates when faced with limited food availability,irrespective of previous experience. The birds harvested seedsfaster when they cached instead of eating seeds; as a result,long-term weight regulation in the poor environment was affectedby caching decisions. Birds tested in late spring lost weightin the poor environment, whereas those tested in the wintermaintained a stable weight when switched from the rich to thepoor environment. Thus, annual endogenous cues directly affectcaching decisions and indirectly influence the long-term regulationof body mass     

The influence of size, shape, and phenolic content on the selection of winter foods by the Azores bullfinch (Pyrrhula murina)

Although the quality of foods may change through the seasons, few studies have assessed shifts in food preferences. I studied winter food selection of the Azores bullfinch to evaluate whether physical characteristics (size and accessibility), or phenolic content of food items better explained changes of winter diet. The birds preferred larger items of all winter foods: seeds, sori and flower buds and more accessible seeds. In autumn, seeds seemed preferred to sori, but in spring birds switched from seeds to sori and flower buds, once the latter became larger. Phenols may influence food preference but they did not explain diet shifts or preferences for individual trees. Food selection is a complex mechanism, depending on the alternatives available and seasonal shifts in food preference may continually change the availability of 'good quality food'.     

Prenatal and early sucking influences on dietary preference in newborn, weaning, and young adult cats

Early experiences are of potential importance in shaping long-term behavior. This study examined the relative influence of prenatal and/or early postnatal experience of chemosensory stimuli on subsequent olfactory and dietary preferences of cats as newborns, at 9-10 weeks, and at 6 months. Cats were exposed to vanillin or 4-ethylguaiacol via their mother's diet either prenatally, postnatally, perinatally (prenatal and postnatal), or experienced no exposure to the stimuli (control). Newborns were given a two-choice olfactory test between the familiar "odor" and no odor; 9-10 week olds were tested for their preference between two food treats, one flavored with the familiar stimulus and the other unflavored; at 6 months, cats were given a choice of two bowls of food, one flavored with the familiar stimulus and the other unflavored. At all ages, cats preferred the familiar, and avoided the unfamiliar, stimulus. Perinatal exposure exerted the strongest influence on preference. Prenatal exposure influenced preference at all ages and postnatal exposure exerted a stronger effect as the cat aged. We conclude that long-term chemosensory and dietary preferences of cats are influenced by prenatal and early (nursing) postnatal experience, supporting a natural and biologically relevant mechanism for the safe transmission of diet from mother to young.     

Food choice by Blue-gray Tanagers in relation to protein content

We tested discriminatory ability and food choice in relation to protein content of the diet in wild-caught Blue-gray Tanagers (Thraupis episcopus), a generalist tropical frugivorous bird. In two sets of experiments we offered to five individual birds in pair-wise choice trials two nearly iso-caloric experimental diets differing in their protein content only. Protein contents of the experimental diets were 4.6 vs. 1.4% in the first experiment and 3.2 and 1.5% (dry matter basis) in the second experiment. Response varied among individual tanagers, but 6 of the 10 birds showed a clear preference for the food highest in protein. Two individuals displayed a strong positional preference. When testing each treatment group, birds ate daily significantly more of the food that had higher protein content. We conclude that Blue-gray Tanagers prefer richer nitrogen foods. Our results also demonstrate that Blue-gray Tanagers have remarkable discriminatory abilities, they reacted to differences in protein content as small as 0.09% fresh matter. We show for the first time discriminatory ability and preference of wild frugivorous birds for foods richer in protein under controlled conditions. Our findings support the hypothesis that frugivorous birds can act as selective agents for fruit pulp composition.     

Innate and Learned Colour Preference in the Zebrafish, Danio rerio

Receiver biases towards specific sensory signals have been demonstrated in insects, birds and fish, both in the context of foraging and mate choice. In some cases, signals important in sexual selection appear to have evolved by exploiting a pre-existing bias in the sensory system. For instance, female preferences for male nuptial colouration may have arisen from selection on foraging practices. Using the zebrafish ( Danio rerio ), a species in which red is not a factor in mate choice, we tested for a foraging bias towards the colour red. We further investigated the plasticity of foraging biases by raising groups of fish on diets consisting solely of red, blue, green or white food. When we subsequently tested their colour preferences in a foraging context, each group responded most strongly to red, irrespective of the colour of food with which they had been conditioned. We also detected a significant effect of conditioning on colour preferences; fish responded more strongly to the colour that matched diet colour than to other colours. The observed receiver bias towards red may have evolved as an adaptive preference for carotenoid compounds in their diet. While the bias to red appears to be innate, our results indicate that learning is also important in shaping foraging biases.     

Food imprinting and subsequent prey preference in the lynx spider,Oxyopes salticus (Araneae: Oxyopidae)

This object of this study was to determine if early feeding experience can affect subsequent prey choice in lynx spiderlings (Oxyopes salticus), and if so, is it evidence for food/olfactory imprinting. After emergence from the egg sac, three groups of 10 spiderlings were each fed exclusively for a 1-week period on one of three naturally-occurring prey species (crickets): group 1 fed on nymphs of Gryllotalpa hexadactyla; group 2 (Dialeurodes citrifoli); group 3 (Microcentrum rhombifolium). Following this, they were tested for subsequent prey preference in choice tests conducted in a plastic arena. Each spiderling was presented simultaneously with one individual of each prey species in a randomized design. Spiderlings exhibited a significant first preference for the original diet. Thus, experience with certain foods encountered by newly hatched spiderlings can affect subsequent prey preference in this species. Given the primacy of the early experience, the results are best interpreted in terms of imprinting and represent the first demonstration of olfactory imprinting in a spider.     

Comparing the Dispersal of Large-seeded Tree Species by Frugivore Assemblagesin Tropical Montane Forest in Africa

We examined frugivore visitation and seed dispersal of five large-seeded (≥ 5 mm) tree species in tropical montane forest based on their occurrence in frugivorous primate diets: Ekebergia capensis, Olea capensis, Parinari excelsa, Prunus africana , and Syzygium guineense. A total of 21 frugivores in five assemblages ( i.e. , chimpanzees, cercopithecines, large-bodied birds, small-bodied birds, and squirrels) were observed over the study period (August 2006 and October–April 2007). We observed seed dispersal in four of five tree species studied; no dispersal was observed for P. excelsa . Frugivore assemblages did not visit tree species equally. Primates spent the most time in trees and had the largest group size. Large-bodied birds (LB) and chimpanzees dispersed the highest number of seeds per minute. LB and cercopithecines potentially dispersed the greatest number of seeds for E. capensis , and chimpanzees for S. guineense . Our analyses indicated that the mean fruiting duration of the focal tree, time in the tree, and number of species present are important predictor variables for seed dispersal by small- and large-bodied birds, and cercopithecines. The number of fruiting trees in the immediate vicinity of the focal tree further predicted seed dispersal for small-bodied birds (SB). Large-bodied birdseed dispersal also was predicted by time in tree by SB, and the number of individuals for SB and cercopithecines. Cercopithecines (CS) were further explained by the time in tree and number of species (SB & LB), and number of individuals for CS. Our study highlights the complexity of describing the relative importance of a frugivore assemblage to the dispersal of a tree species seeds.     

Food preference of young rhesus monkeys fed marginally zinc deficient diets

Zinc deprivation has been shown to produce hypogeusia in rhesus monkeys but the possible consequences of altered taste sensitivity for food preference and selection is not known. We studied 13 rhesus monkeys (2.5 to 3 years of age) for food preference in a structured choice situation. These animals were fed two levels of dietary zinc: marginally zinc deprived (fed 4 μg zinc/g diet from conception and throughout development;N=6), or control (fed a 100 μg zinc/g diet throughout the study;N=7). Preference for familiar vs unfamiliar food items, order of food preferences, persistence in retrieval of preferred and nonpreferred foods, and preference under four deprivation periods were examined. Animals were offered a choice of two food items, both containing minimal zinc levels, and food choice and latency of choice were measured. Results indicate that animals fed the marginal zinc diets had reduced preference for unfamiliar foods relative to controls and different patterns of food preference. These results may be relevant to maintenance of appropriate food selection in marginally zinc deprived populations.     

Effects of bird ingestion on seed germination of Sorbus commixta

To determine the effects of ingestion by birds on seed germination, we performed germination experiments in the field and laboratory with Sorbus commixta. The germination of four groups of seeds was compared: ingested seeds, seeds defecated in feces after feeding of fruits to birds; extracted seeds, seeds deliberately extracted from the fruit pulp; juiced seeds, seeds plus the juice of the pulp after seeds had been deliberately extracted from the pulp; intact seeds, seeds in untreated intact fruits. In the laboratory, intact and juiced seeds hardly germinated, but ingested and extracted seeds germinated. Thus, the pulp and its juice appeared to inhibit germination, but seeds could germinate without ingestion by birds once the seeds had been manually extracted from the pulp. In the field, intact fruits did not germinate in the first spring, because the seed was still covered with pulp. The pulp of intact seeds decomposed during the first summer, and thus, the seeds had the potential to germinate during the second spring. In fact, most intact seeds do not germinate during the second spring either, since they lose their viability during the first summer. Thus, under natural conditions, most seeds of Sorbus commixta cannot germinate without bird ingestion. Received: 5 July 1997 / Accepted: 7 November 1997     

The Influence of Prior Learning Experience on Pollinator Choice: An Experiment Using Bumblebees on Two Wild Floral Types of Antirrhinum majus

Understanding how pollinator behavior may influence pollen transmission across floral types is a major challenge, as pollinator decision depends on a complex range of environmental cues and prior experience. Here we report an experiment using the plant Antirrhinum majus and the bumblebee Bombus terrestris to investigate how prior learning experience may affect pollinator preferences between floral types when these are presented together. We trained naive bumblebees to forage freely on flowering individuals of either A. majus pseudomajus (magenta flowers) or A. majus striatum (yellow flowers) in a flight cage. We then used a Y-maze device to expose trained bumblebees to a dual choice between the floral types. We tested the influence of training on their choice, depending on the type of plant signals available (visual signals, olfactory signals, or both). Bumblebees had no innate preference for either subspecies. Bumblebees trained on the yellow-flowered subspecies later preferred the yellow type, even when only visual or only olfactory signals were available, and their preference was not reinforced when both signal types were available. In contrast, bumblebees trained on the magenta-flowered subspecies showed no further preference between floral types and took slightly more time to make their choice. Since pollinator constancy has been observed in wild populations of A. majus with mixed floral types, we suggest that such constancy likely relies on short-term memory rather than acquired preference through long-term memory induced by prior learning.     

The influence of substrate on the functional response of an avian granivore and its implications for farmland bird conservation

Few studies to date have considered the effect of substrate on the functional response of an organism feeding on prey of varying visibility. Intake rates of lone captive canaries, Serinus canarius L., were measured at varying seed densities on patches of either earth or short grass (<1 cm). Experiment 1, using pale seeds, found intake rates were significantly higher and search times significantly lower on earth than on grass. Two measures of crypticity (contrast in light reflectance as measured using a spectrophotometer and an experiment with humans) found pale seeds to be more visible on earth. The results from experiment 1 could be explained by this difference in crypsis. Experiment 2 used identical seeds to those in experiment 1 except they were dyed to match their backgrounds. The two measures of crypticity both found that black seeds were less visible on earth than green seeds were on grass. However, intake rates were still significantly higher on earth than grass. Seed colour preference, vegetation impeding movement, and differences in vigilance rates or seed accessibility could not explain this result. We discuss three other potentially explanatory mechanisms, the most likely of which was the greater surface area needed for scanning created by the structure of grass. Crucially, regardless of the mechanism(s) involved, many vegetated substrates share similar properties with grass (structural complexity and shiny surfaces which reflect light) and so the outcome of our findings are likely to extend to many natural situations. Conservationists wishing to encourage granivorous birds should consider enhancing food accessibility by providing uniform substrates, such as bare earth, for them to forage on. In addition, behaviour-based models should incorporate the effects of habitat into their equations of the functional response.     

Does natal habitat preference modulate cereal kernel preferences in the rice weevil?

The influence of larval diet source in Sitophilus oryzae (L.), a primary grain feeder, on the choice of a new cereal host by young adult was investigated through a multiple choice comparative bioassay. For this purpose, virgin male and female adults, which had been reared as immatures in different grain commodities, were monitored according to their food preference in cylindrical arenas containing whole kernels of maize, rice, barley and wheat. The adults showed a clear individual preference for maize kernels, regardless of the previous food experience, sex and also the interval on which the adults are exposed to food. Males were much more mobile than females and visited more vigorously different commodities before the final choice. Female response was found to be related with male response to specific food sources. Hence, our results indicate that there was no natal habitat preference induction, since maize was, in most cases, the final commodity choice.     

Macrogeographic variations in food choice of mainland citril finches Carduelis [citrinella] citrinella versus insular Corsican (citril) finches Carduelis [citrinella] corsicanus

We studied variations in diet composition in sub-populations of mainland citril finches Carduelis [citrinella] citrinella in European mountain chains, compared to closely related insular Corsican (citril) finches Carduelis [citrinella] corsicanus on several Mediterranean Islands with a special emphasis on key food plants used during the breeding period. We found that citril finch sub-populations of the Pre-Pyrenees and the Black Forest rely mostly on mountain pine and dandelion seeds as main food resources during the breeding season. In contrast, insular Corsican finches on the Mediterranean islands of Capraia and Corsica fed mainly on various herb species. Finches in Corsica fed predominantly on shepherd’s purse, while birds from Capraia were choosing rosemary as their main food plant during the breeding period. These observations confirmed the previously expected difference in food choice between the two forms due to the observed increased niche breadth on Corsica. However, Corsican finches on Sardinia showed a food choice more similar to that of the citril finch populations, in that they fed predominantly on black pine seeds during breeding season. Thus we were not able to find clear behavioural differences in diet composition between all studied sub-populations of citril and Corsican finches. To conclude, food choice of both forms is highly adaptive and, in the different breeding areas, distinct key plants may be of special importance during the breeding season. The preference for these plants is likely to be linked to their abundance and their energetic and nutritional content.     

Influence of method of whole-wheat feeding on the performance,digestive tract development and carcass traits of broiler chickens

The present study was conducted to investigate the effects of feeding whole wheat either through a mixed feeding (MF) or free choice feeding (FCF) system on the performance, digestive tract development and carcass traits of broiler chickens. The following three treatments, based on wheat and soybean meal, were employed: GW, ground-wheat diet with 600–690 g wheat kg⁻¹; MF, GW diet with 490–500 g wheat kg⁻¹ and 100–200 g whole wheat kg⁻¹; and FCF, whole wheat and a protein concentrate offered in separate feeders. Each diet was fed to six pens of 36 birds each from day 7 to 35 post-hatch. Over the 7–35-day trial period, no differences (P>0.05) were observed between the weight gain, feed intake and feed per gain of broilers receiving the GW and MF treatments. Birds receiving the FCF treatment had the lowest (P<0.05) weight gain and feed intake, and the highest (P<0.05) feed per gain. During week 1 of the trial, the protein concentrate was consumed more than the whole wheat (0.69 vs. 0.31) in the FCF treatment, resulting in the amount of concentrate offered being restricted during subsequent weeks. Over the trial period, the average consumption of protein concentrate remained high (0.56 of the total intake). Both whole-wheat treatments increased (P<0.05) the relative gizzard weights. Factors that affect diet selection such as learning and previous experience, visual differences between the foods, texture and flavour of the food, and palatability may explain the lower whole-wheat intake in the FCF treatment. In this study, whole wheat was introduced only on day 7 and it is possible that the birds may have to be trained to experience choice feeding from the first week of life. However, the present results suggest that FCF may not be an appropriate feeding system for fast growing modern broilers.