Foraging preferences: Response to reward skew

Recent theoretical and empirical work on foraging behaviour suggests that animals may respond to both the means and variances in benefits associated with available resources. We attempt to extend this analysis by asking if reward skew (third moment about the mean) might influence preference when two options have equal means and equal variances. We examine how minimizing the probability of starvation might induce response to skew. In the Appendix we develop an expected ‘fitness’ model which follows from economic theory and indicates more general conditions concerning responses to skew. We also report experiments involving foraging white-crowned sparrows (Zonotrichia leucophrys). Under conditions where positive skew should be favoured, the birds' behaviour supports the prediction. However, their response to skew is not as strong as responses to variance noted in the same individuals.     

Song development in the white-crowned sparrow: modification of learned song

Previous research has shown that laboratory-tutored white-crowned sparrows, Zonotrichia leucophrys nuttalli, can learn tape-recorded song only until they are about 50 days of age. However, they can learn either conspecific or allospecific song after they are 50 days of age when a live tutor is used. Because the live-tutored birds had not been exposed to song during the first 50 days of life, a question regarding the reasonableness of applying these results to the natural, field situation can be raised. In the experiments reported here, hand-raised nestling white-crowned sparrows between 10 and 50 days of age were tutored in the laboratory with one song, and were then placed with a live tutor singing a different dialect when they were over 50 days of age. Eight of 13 males, and none of 10 females, adopted the song of the second tutor. Birds were also captured in the field as fledglings, and were placed with a live tutor singing a song different from that of the natal area: three of seven males, and one of five females, adopted the song of the live tutor. Four birds that had been group-isolated (two males and two females) were exposed to a live tutor when they were 100 days old, and none acquired normal song; all sang isolate song. The results of these studies indicate that there is considerable plasticity in the song learning of whitecrowned sparrows, and that, for many individuals, song can be modified quite readily. The implications of these findings for the nature of the sensitive phase and for mechanisms of song learning are discussed.     

One meadow for two sparrows: resource partitioning in a high elevation habitat

Resource partitioning is the basis of the coexistence of sympatric species and has therefore received much attention in ecological studies. However, how variation in environmental conditions (and particularly natural variation in resource availability) can influence resource partitioning in free-ranging animals is not well understood. In the present study, we addressed the hypothesis that natural changes in the availability of food resources affect food partitioning between sympatric species. To do so, we examined temporal changes in the plasma isotopic signature (δ(15)N and δ(13)C) of syntopic Lincoln's sparrows Melospiza lincolnii and white-crowned sparrows Zonotrichia leucophrys, in parallel with seasonal changes in habitat maturity and food availability from spring to early summer. We found no apparent trophic segregation between Lincoln's and white-crowned sparrows when resources were scarce in spring. But, interestingly, as resource availability and the number of breeding birds increased, Lincoln's sparrows showed lower δ(15)N values than white-crowned sparrows, as they consumed more prey from lower trophic levels and less prey from higher trophic levels. This feeding divergence between sympatric species may be explained (1) by a change in foraging preferences and opportunities for Lincoln's sparrows and (2) by the abundance of competitors that increased faster than resources, thus promoting interspecific competition and trophic segregation. These results provide clear evidence that trophic segregation is dynamically tied to variation in environmental conditions, which are therefore fundamental to consider when examining resource partitioning between co-existing species.     

Assortative mating by white-crowned sparrows at song dialect boundaries

We tested the hypothesis that female Nuttall's white-crowned sparrows (Zonotrichia leucophrys nuttalli) mate assortatively with males from the same dialect population. Young birds of both sexes learn their natal dialect during an early sensitive period, and for females, this early experience may be the basis of future mate choice. A total of 32 female sparrows were mist-netted for three experiments at or near dialect boundaries just inside the Limantour, Drake and Clear dialect populations in Marin Co., California. The mates of all captured females sang the home dialect. All females were implanted with testosterone and maintained in captivity. Of the females which began to sing, 25 sang recognizable white-crowned sparrow songs. Of those 25 females, 24 had mated assortatively, thus supporting the assortative mating hypothesis.     

Signals of status in wintering white-crowned sparrows, Zonotrichia leucophrys gambelii

The possibility of plumage status signalling within the social systems of wintering birds has been a controversial issue. Our results are the first to demonstrate conclusively the reality of such signalling. Data from eight groups of captive white-crowned sparrows (Zonotrichia leucophrys gambelii), each with 8 to 11 different individuals, show that immature and adult females with crowns painted to resemble more brightly coloured, dominant adult males consistently win encounters with control birds of their own age and sex. These experiments demonstrate that signals that correlate with age (adult versus immature) and sex (adult male versus adult female) are used by the birds as reliable indicators of relative dominance position. Our demonstration of status signalling draws attention to the need to explain how such a system can be evolutionarily stable and we discuss some suitable models.     

Proportions of mammalian-type and reptilian-type nephrons in the kidneys of two passerine birds

We compared the proportions of mammalian-type and reptilian-type nephrons in the kidneys of two species of passerine birds. The desert house sparrow (Passer domesticus) is relatively well adapted for water conservation, whereas the white-crowned sparrow (Zonotrichia leucophrys) is more mesic adapted. The two species do not differ in body mass, but the kidneys of P. domesticus are significantly smaller than those of Z. leucophrys. Associated with its smaller size, the house sparrow kidney has significantly fewer glomeruli (35,700 per kidney) than does the white-crowned sparrow kidney (53,000 per kidney). The medullary cones, which contain the loops of Henle of the mammalian-type nephrons, are significantly longer in house sparrows than in white-crowned sparrows (2.2 vs. 1.9 mm). The number of medullary cones, the number of nephrons per medullary cone, and, hence, the number of mammalian-type nephrons do not differ between the two species. The smaller number of nephrons in the kidney of the house sparrow therefore represents a smaller number of reptilian-type nephrons. Desert house sparrows have 18% mammalian-type nephrons, whereas white-crowned sparrows have 10% mammaliantype nephrons. The relative reduction of reptilian-type nephrons in P. domesticus may reduce the flow of dilute urine through the collecting ducts, thereby permitting a greater concentration gradient to be established along the length of the medullary cones.     

Rationality in risk-sensitive foraging choices by starlings

Normative models of choice usually predict preferences between alternatives by computing their value according to some criterion, then identifying the alternative with greatest value. An important consequence of this procedure is captured in the economic concept of rationality, defined through a number of principles that are necessary for the existence of an ordinal scale of value upon which organisms base their choices. Violations of these principles, such as some recently reported breaches of transitivity and regularity in birds and honeybees, have strong implications for the understanding of decision mechanisms in humans and nonhumans alike. We investigated rationality in risky choice using European starlings, Sturnus vulgaris. Birds had to choose between two or three food sources, each associated with a different variance in delay to reward. In three experiments, starlings were strongly risk prone, showing regular and consistent preferences in binary and trinary choices. Preferences also satisfied weak and strong stochastic transitivity. Our results extend the generality of previous research in risk-sensitive foraging to situations where more than two alternatives are present and suggest that violations of rationality in risk-sensitive choices may be expressed only under restricted sets of conditions. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Hippocampal neurogenesis is associated with migratory behaviour in adult but not juvenile sparrows (Zonotrichia leucophrys ssp.)

It has been hypothesized that individuals who have higher demands for spatially based behaviours should show increases in hippocampal attributes. Some avian species have been shown to use a spatially based representation of their environment during migration. Further, differences in hippocampal attributes have been shown between migratory and non-migratory subspecies as well as between individuals with and without migratory experience (juveniles versus adults). We tested whether migratory behaviour might also be associated with increased hippocampal neurogenesis, and whether potential differences track previously reported differences in hippocampal attributes between a migratory (Zonotrichia leucophrys gambelii) and non-migratory subspecies (Z. l. nuttalli) of white-crowned sparrows. We found that non-migratory adults had relatively fewer numbers of immature hippocampal neurons than adult migratory birds, while adult non-migrants had a lower density of new hippocampal neurons than adult and juvenile migratory birds and juvenile non-migratory birds. Our results suggest that neurogenesis decreases with age, as juveniles, regardless of migratory status, exhibit similar and higher levels of neurogenesis than non-migratory adults. However, our results also suggest that adult migrants may either seasonally increase or maintain neurogenesis levels comparable to those found in juveniles. Our results thus suggest that migratory behaviour in adults is associated with maintained or increased neurogenesis and the differential production of new neurons may be the mechanism underpinning changes in the hippocampal architecture between adult migratory and non-migratory birds.     

Avian orientation: effects of cue-conflict experiments with young migratory songbirds in the high Arctic

The migratory orientation of juvenile white-crowned sparrows, Zonotrichia leucophrys gambelli, was investigated by orientation cage experiments in manipulated magnetic fields performed during the evening twilight period in northwestern Canada in autumn. We did the experiments under natural clear skies in three magnetic treatments: (1) in the local geomagnetic field; (2) in a deflected magnetic field (mN shifted −90°); and (3) after exposure to a deflected magnetic field (mN −90°) for 1 h before the cage experiment performed in the local geomagnetic field at dusk. Subjects showed a mean orientation towards geographical east in the local geomagnetic field, north of the expected migratory direction towards southeast. The sparrows responded consistently to the shifted magnetic field, demonstrating the use of a magnetic compass during their first autumn migration. Birds exposed to a cue conflict for 1 h on the same day before the experiment, and tested in the local geomagnetic field at sunset, showed the same northerly orientation as birds exposed to a shifted magnetic field during the experiment. This result indicates that information transfer occurred between magnetic and celestial cues. Thus, the birds' orientation shifted relative to available sunset and geomagnetic cues during the experimental hour. The mean orientation of birds exposed to deflected magnetic fields prior to and during testing was recorded up to two more times in the local geomagnetic field under natural clear and overcast skies before release, resulting in scattered mean orientations.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Cues for Early Discrimination of Conspecific Song in the White-Crowned Sparrow (Zonotrichia leucophrys)

In song learning, white-crowned sparrows ( Zonotrichia leucophrys ) begin memorizing conspecific song models at around 20 d of age. Even prior to song memorization, however, between 10 and 20 d of age, these birds respond differently to playbacks of conspecific and heterospecific songs. To investigate the acoustic cues underlying this early song discrimination, we measured the vocal responses of newly fledged young to playbacks of modified conspecific and heterospecific songs. Fledgling white-crowned sparrows responded more strongly to songs containing conspecific notes than to songs containing notes from other species. In contrast, the presence or placement of an introductory whistle, present in all white-crowned sparrow songs, did not affect response levels. A whistle has previously been shown to serve as an acoustic cue for song memorization and production in this species. Such different responses to the species-typical introductory whistle at different stages suggests that developmental changes occur in the template system underlying song learning, the factors governing song recognition, or both.     

Photorefractoriness in a sparrow: Phase of circadian photosensitivity elucidated by skeleton photoperiods

Summary Photorefractoriness can be maintained in first-year male white-crowned sparrows kept on short daylengths if the birds are exposed to a short light pulse interrupting the dark period during a daily phase of photosensitivity. The photosensitive phase appears to be located between hours 9 and 20 following the onset of first light (Fig. 1).     

The effect of corticosterone on standard metabolic rates of small passerine birds

Summary Resting metabolic rates of Gambel's white-crowned sparrows (Zonotrichia leucophrys gambelii) and pine siskins (Carduelis pinus) were evaluated at thermoneutral temperatures before and after administration of corticosterone (B) at physiological doses. There was no effect of B on basal metabolic rate of either species, but nocturnal metabolic rate varied significantly less over the 3-h period of measurement in B-treated sparrows and siskins than in control birds. These results, coupled with observations of caged birds, suggest that corticosterone has no direct effect on avian resting metabolism but does reduce the responsiveness of birds to external stimuli and thus promotes nocturnal restfulness.Abbreviations B corticosterone - VO ₂ rate of oxygen consumption - T ₃ tri-iodothyromine - T ₄ tetra-iodothyronine, thyroxine     

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.     

Migratory sleeplessness in the white-crowned sparrow (Zonotrichia leucophrys gambelii)

Twice a year, normally diurnal songbirds engage in long-distance nocturnal migrations between their wintering and breeding grounds. If and how songbirds sleep during these periods of increased activity has remained a mystery. We used a combination of electrophysiological recording and neurobehavioral testing to characterize seasonal changes in sleep and cognition in captive white-crowned sparrows (Zonotrichia leucophrys gambelii) across nonmigratory and migratory seasons. Compared to sparrows in a nonmigratory state, migratory sparrows spent approximately two-thirds less time sleeping. Despite reducing sleep during migration, accuracy and responding on a repeated-acquisition task remained at a high level in sparrows in a migratory state. This resistance to sleep loss during the prolonged migratory season is in direct contrast to the decline in accuracy and responding observed following as little as one night of experimenter-induced sleep restriction in the same birds during the nonmigratory season. Our results suggest that despite being adversely affected by sleep loss during the nonmigratory season, songbirds exhibit an unprecedented capacity to reduce sleep during migration for long periods of time without associated deficits in cognitive function. Understanding the mechanisms that mediate migratory sleeplessness may provide insights into the etiology of changes in sleep and behavior in seasonal mood disorders, as well as into the functions of sleep itself.     

The relationship between uric acid and its oxidative product allantoin: a potential indicator for the evaluation of oxidative stress in birds

Uric acid is the main nitrogenous waste product in birds but it is also known to be a potent antioxidant. Hominoid primates and birds lack the enzyme urate oxidase, which oxidizes uric acid to allantoin. Consequently, the presence of allantoin in their plasma results from non-enzymatic oxidation. In humans, the allantoin to uric acid ratio in plasma increases during oxidative stress, thus this ratio has been suggested to be an in vivo marker for oxidative stress in humans. We measured the concentrations of uric acid and allantoin in the plasma and ureteral urine of white-crowned sparrows (Zonotrichia leucophrys gambelii) at rest, immediately after 30 min of exercise in a hop/hover wheel, and after 1 h of recovery. The plasma allantoin concentration and the allantoin to uric acid ratio did not increase during exercise but we found a positive relationship between the concentrations of uric acid and allantoin in the plasma and in the ureteral urine in the three activity phases. In the plasma, the slope of the regression describing the above positive relationships was significantly higher immediately after activity. We suggest that the slope indicates the rate of uric acid oxidation and that during activity this rate increases as a result of higher production of free radicals. The present study demonstrates that allantoin is present in the plasma and in the ureteral urine of white-crowned sparrows and therefore might be useful as an indicator of oxidative stress in birds.     

Pink flower preference in sunbirds does not translate into plant fitness differences in a polymorphic Erica species

Bird-pollinated plants typically have reddish flowers, but it is not clear whether this trait can be attributed to selection by birds. Here we experimentally test for the first time the foraging behaviour of sunbirds in relation to flower colour, using the Orange-breasted Sunbird Anthobaphes violacea (Nectariniidae) and the colour dimorphic Erica perspicua (Ericaceae). Pink and white flower morphotypes co-flower in intermixed populations and have similar nectar volumes and concentrations. Using floral arrays in a field aviary, we found that sunbirds preferred pink flowers; 95 % of their first choices were to pink inflorescences and they visited and probed more pink inflorescences and flowers, respectively. We also tested for flower constancy (the tendency to move between same colour rather than different colour morphotypes), but found no evidence for this in the sequence of their foraging choices, indicating that this mechanism did not maintain flower colour differences in sympatry. There was evidence for optimal foraging: 80 % of moves were to adjacent inflorescences. Unexpectedly, the preference for pink flowers observed in the aviary did not translate into a female fitness advantage for this morphotype in the field, since no difference is found in natural pollination rate, fruit or seed set. This may be because the minimization of flight distances between plants is the primary factor in sunbird foraging choices, overriding their colour preference. Antagonistic nectar robbers did not act as a selective force on the polymorphism, since nectar-robbing rates were equal between white and pink morphotypes in the field.     

Evidence for androgen independence of male mounting behavior in white-crowned sparrows (Zonotrichia leucophrys gambelii)

Previous experiments have shown that expression of mounting behavior in sexually inexperienced, adult male white-crowned sparrows does not require elevated plasma levels of androgen; adult males maintained on nonstimulatory short days mount sexually receptive females. The experiments reported here demonstrate that (1) sexually inexperienced, prepubertal males maintained on nonstimulatory short days show very low mounting rates in response to female sexual displays; (2) these males exhibit high mounting rates when exposed to stimulatory long days but androgen treatment on short days is ineffective in stimulating high mounting rates; and (3) prepubertal castration has no effect on the expression of mounting behavior by photostimulated adult males. Thus, there is no evidence that mounting behavior of adult male white-crowned sparrows depends on androgen.     

Learning to Choose Among Social Foraging Strategies in Adult House Sparrows (Passer domesticus)

Social foragers may be regarded as being engaged in a producer–scrounger game in which they can search for food independently or join others who have discovered food. Research on the producer–scrounger game has focused mainly on the different factors influencing its evolutionarily stable strategy (ESS) solution, but very little is known about the actual mechanisms that shape players' decisions. Recent work has shown that early experience can affect producer–scrounger foraging tendencies in young house sparrows and that in nutmeg mannikins learning is involved in reaching the ESS. Here, we show that direct manipulation of the success rate experienced by adult sparrows when following others can change their strategy choice on the following day. We presented to live sparrows an experimental regime, where stuffed adult house sparrows in a feeding position were positioned on a foraging grid that included two reward regimes: a positive one, in which the stuffed models were placed near food, and a negative one, in which the models were placed away from food. There was a significant increase in joining behavior after the positive treatment (exhibited by 84% of the birds), but no change after the negative treatment. Further analysis demonstrated that sparrows more frequently used the strategy with which they were more successful (usually joining) and that differences in strategy use were correlated with differences in success. These results suggest that adult birds can monitor their success and learn to choose among social foraging strategies in the producer–scrounger game.