A core assumption implicit in economic models of animal choice is that subjects assign absolute utilities to options that are independent of the type and number of alternatives available. Humans sometimes appear to violate this assumption and employ relative, as opposed to absolute, currencies when making choices. Recent evidence suggests that animals too might sometimes employ relative choice mechanisms. We tested this idea by measuring the foraging preferences of rufous hummingbirds (Selasphorus rufus) faced with choices analogous to those in which human use of relative currencies is evident. The birds experienced three treatments: a binary choice between two artificial flower types designated concentration (20 microl, 40% sucrose solution) and volume (40 microl, 20%), and two trinary treatments in which a third decoy option (either concentration decoy: 10 microl, 30% or volume decoy: 30 microl, 10%) was added to the set. The birds' preferences differed significantly across the three treatments. In the trinary treatments, the effect of the decoy options was to increase the preference for the option that dominated the decoy. These results are similar to those reported in the human choice literature, and are compatible with the hummingbirds using a relative evaluation mechanism in decision making.
Animals organize their lives around circannual and circadian rhythms, but little is known of their use of much shorter intervals. In the laboratory, some animals can learn the specific duration (seconds or minutes) between periods of food access. It has been supposed that wild nectarivores, such as hummingbirds, might also learn short time intervals so as to avoid revisiting emptied flowers until the nectar has been replenished. We provided free-living, territorial rufous hummingbirds each with eight artificial flowers containing sucrose solution. Four flowers were refilled 10 min after the bird emptied them, and the other four were refilled 20 min after being emptied. Throughout the day, birds revisited the 10 min flowers significantly sooner than they revisited the 20 min flowers, and return visits to the flowers matched their refill schedules. Hummingbirds remembered the locations and timing of eight rewards, updating this information throughout the day. Not only is this the first time that this degree of timing ability has been shown in wild animals, but these hummingbirds also exhibit two of the fundamental aspects of episodic-like memory (where and when), the kind of memory for specific events often thought to be exclusive to humans.
An animal's behavior is affected by its cognitive abilities, which are, in turn, a consequence of the environment in which an animal has evolved and developed. Although behavioral ecologists have been studying animals in their natural environment for several decades, over much the same period animal cognition has been studied almost exclusively in the laboratory. Traditionally, the study of animal cognition has been based on well-established paradigms used to investigate well-defined cognitive processes. This allows identification of what animals can do, but may not, however, always reflect what animals actually do in the wild. As both ecologists and some psychologists increasingly try to explain behaviors observable only in wild animals, we review the different motivations and methodologies used to study cognition in the wild and identify some of the challenges that accompany the combination of a naturalistic approach together with typical psychological testing paradigms. We think that studying animal cognition in the wild is likely to be most productive when the questions addressed correspond to the species' ecology and when laboratory cognitive tests are appropriately adapted for use in the field. Furthermore, recent methodological and technological advances will likely allow significant expansion of the species and questions that can be addressed in the wild.
Most hermaphroditic, many‐flowered plants should suffer reduced fitness from within‐plant selfing (geitonogamy). Large inflorescences are most attractive to pollinators, but also promote many flower visits during a single plant visit, which may increase selfing and decrease pollen export. A plant might avoid the negative consequences of attractiveness through modification of the floral display to promote fewer flower visits, while retaining attractiveness. This report shows that increasing only the variance in nectar volume per flower results in fewer flower visits per inflorescence by wild hummingbirds (Selasphorus rufus) and captive bumble bees (Bombus flavifrons) foraging on artificial inflorescences. Inflorescences were either constant (all flowers contained the same nectar volume) or variable (half the flowers were empty, the other half contained twice as much nectar as in the constant flowers). Both types of inflorescence were simultaneously available to foragers. Risk‐averse foraging behaviour was expressed as a patch departure preference: birds and bees visited fewer flowers on variable inflorescences, and this preference was expressed when resource variability could be determined only by concurrent sampling. When variance treatments were clearly labelled with colour and offered to hummingbirds, the departure effect was maintained; however, when preference was measured by inflorescence choice, birds did not consistently prefer to visit constant inflorescences. The reduced visitation lengths on variable inflorescences by both birds and bees documented in this study imply that variance in nectar production rates within inflorescences may represent an adaptive trait to avoid the costs of geitonogamy.
The spatial memory abilities of free-ranging rufous hummingbirds were tested in an "open-field" analogue of a radial maze. Eight artificial "flowers" filled with sucrose solution and arranged in a circle were placed within each bird's feeding territory. The birds were given two types of trial: in "free" trials, birds could choose which four of eight flowers to feed from before leaving the array; in "forced" trials, the experimenter presented birds with four flowers. Memory performance was assessed by observing the ability of the birds to avoid the emptied flowers on the return visit. On both types of trials, hummingbirds were able to avoid visiting the previously emptied flowers for retention intervals ranging from a few minutes to over an hour. Demonstration of good memory performance in free-ranging, nondeprived animals has both ethical and logistical implications for future studies on animal memory. In addition, such studies offer an ideal opportunity to test what and how well animals remember in their natural environment.
Black-capped Chickadee (Parus atricapillus) song consists of two notes, termed fee and bee. Frequency measures at three key points (at the start and end offee, and at the start of bee) were obtained from the songs of a large sample of chickadees (n = 15 1) in the wild. In this sample, 19 birds produced songs shifted downward in frequency as well as their normal songs. Analysis of normal song revealed that fee declines in frequency in a glissando of nearly pure tone, then continues at greatly reduced amplitude at the start of bee; whereas bee, also a nearly pure tone, is always lower in frequency than either the start or end offee. The absolute pitches (frequencies) of these measures vary substantially among birds, but much less within individuals. In contrast, pitch intervals (ratios of higher to lower frequencies) for frequency changes among the three measures are highly invariant among birds. Moreover, chickadees with normal and frequency-shifted songs maintain virtually the same pitch intervals in both. This analysis suggests that the absolute and relative pitch constancies in chickadee song production may provide information for individual and species recognition, respectively.
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