Win-shift and win-stay learning in the short-beaked echidna (<Emphasis Type="Italic">Tachyglossus aculeatus</Emphasis>)

Numerous previous investigators have explained species differences in spatial memory performance in terms of differences in foraging ecology. In three experiments we attempted to extend these findings by examining the extent to which the spatial memory performance of echidnas (or "spiny anteaters") can be understood in terms of the spatio-temporal distribution of their prey (ants and termites). This is a species and a foraging situation that have not been examined in this way before. Echidnas were better able to learn to avoid a previously rewarding location (to "win-shift") than to learn to return to a previously rewarding location (to "win-stay"), at short retention intervals, but were unable to learn either of these strategies at retention intervals of 90 min. The short retention interval results support the ecological hypothesis, but the long retention interval results do not. Electronic Publication     

Learning and foraging efficiency in German cockroaches, Blattella germanica (L.) (Insecta: Dictyoptera)

We analysed, under laboratory test conditions, how German cockroach larvae oriented their outgoing foraging trip from their shelter. Our results stressed the importance of external factors, like availability and spatial distribution of food sources, in the choice of a foraging strategy within their home range. When food sources were randomly distributed, larvae adopted a random food search strategy. When food distribution was spatially predictable and reliable, cockroaches were able to relate the presence of food with a landmark during a 3-day training period and to develop an oriented search strategy. Cockroaches were able to associate learned spatial information about their home range to the presence of food resources and then to improve their foraging efficiency. However, conflict experiments revealed that detection of food odour overrode learned landmark cues. Received: 16 October 1999 / Accepted after revision: 18 July 2000     

“To eat or not to be eaten?” Collective risk-monitoring in groups

The importance of risk-monitoring has been increasing in many key aspects of our modern lives. This paper examines how individuals monitor such risks collectively by extending a behavioral ecological model of animal foraging to human groups. Just as animals must forage for food under predatory risk, humans must divide valuable material and psychological resources between foraging activity and risk-monitoring activity. We predicted that game-theoretic aspects of the group situation complicate such a trade-off decision in resource allocation, eventually yielding a mixed equilibrium in a group. When the equilibrium is reached, only a subset of members engage in the risk-monitoring activity while others free-ride, concentrating mainly on their own foraging activity. Laboratory groups engaging in foraging under moderate risk provided a support to this prediction. When the risk-level was set higher, however, “herding behavior” (conforming to the dominant behavior) interfered with the emergence of equilibrium. Implications for risk management are discussed.     

What wild primates know about resources: opening up the black box

We present the theoretical and practical difficulties of inferring the cognitive processes involved in spatial movement decisions of primates and other animals based on studies of their foraging behavior in the wild. Because the possible cognitive processes involved in foraging are not known a priori for a given species, some observed spatial movements could be consistent with a large number of processes ranging from simple undirected search processes to strategic goal-oriented travel. Two basic approaches can help to reveal the cognitive processes: (1) experiments designed to test specific mechanisms; (2) comparison of observed movements with predicted ones based on models of hypothesized foraging modes (ideally, quantitative ones). We describe how these two approaches have been applied to evidence for spatial knowledge of resources in primates, and for various hypothesized goals of spatial decisions in primates, reviewing what is now established. We conclude with a synthesis emphasizing what kinds of spatial movement data on unmanipulated primate populations in the wild are most useful in deciphering goal-oriented processes from random processes. Basic to all of these is an estimate of the animal’s ability to detect resources during search. Given knowledge of the animal’s detection ability, there are several observable patterns of resource use incompatible with a pure search process. These patterns include increasing movement speed when approaching versus leaving a resource, increasingly directed movement toward more valuable resources, and directed travel to distant resources from many starting locations. Thus, it should be possible to assess and compare spatial cognition across a variety of primate species and thus trace its ecological and evolutionary correlates. This contribution is part of the special issue “A Socioecological Perspective on Primate Cognition” (Cunningham and Janson 2007b)     

Chunking versus foraging search patterns by rats in the hierarchically baited radial maze

Rats were exposed to a radial maze containing six black smooth arms and six wire-grid-covered arms and a striped 'exit arm' in experiment 1. The probability of a black or grid arm being baited (5/6 vs 1/6) with sunflower seeds was associated with its proximal cue for some rats (the Relevant Arm Cue group) but not for others (the Irrelevant Arm Cue group). All rats could terminate a trial and receive a highly preferred morsel of apple by entering the exit arm only after having sampled all six seed-baited arms. Relevant Arm Cue rats usually chose some arms from the more densely baited set before choosing an arm from the less densely baited set and made fewer reentries than Irrelevant Arm Cue rats. Although such clustered, higher choice accuracy in the Relevant Arm Cue group corresponds to human clustered, better recall of verbal items from lists hierarchically organized by categories, these rats did not similarly exhaustively retrieve items (arm locations). That is, when required to terminate a trial by entering the 'exit' arm for an apple morsel after having sampled all seed-baited arms, both groups were equally unable to withhold making nonrewarded premature exits. This nonexhaustive foraging search pattern was maintained in the next two experiments in which the radial maze was reduced to three black and three grid arms along with the striped 'exit' arm and in which black and grid arm cues were paired with number of seeds (eight or one) in an arm for Relevant Arm Cue rats. Although Relevant Arm Cue rats displayed perfect clustering by entering all eight-seeded arms before a one-seeded arm, they made more premature exits and reentries into eight-seeded arms in experiment 2 or when forced to enter all eight-seeded arms in experiment 3 than did Irrelevant Arm Cue rats. These foraging tendencies prevent accurate estimations of the amount of information (i.e., arm locations) rats can 'chunk'. Electronic Publication     

Motivation and memory in zebra finch (<Emphasis Type="Italic">Taeniopygia guttata</Emphasis>) foraging behavior

Remembering combinations of information such as what resources have been seen in which locations could play an important role in enhancing individual survival through increased foraging success. To date, there have been relatively few investigations of avian memory involving more than one category of information. This study explored zebra finches’ (Taeniopygia guttata) capacity to recall two categories in combination, namely food-type and spatial location. Birds were trained to remove variously weighted flaps to find two types of food hidden beneath. Memory for food-types and locations was assessed by pre-feeding the birds to satiety on and devaluing one food-type, and then testing the birds’ efficiency at finding the non-devalued food. When allowed one trial to learn locations of two food rewards that were hidden beneath lightly weighted flaps, birds performed better than chance at locating a food reward. However, they did not preferentially search for the non pre-fed food, suggesting that they were unable to recall both food type and location in combination. Zebra finches made fewer errors when tested on the one-trial task using more heavily weighted flaps than with lightly weighted flaps; there was equivocal evidence that they remembered which food type was hidden where on this task. When given repeated exposures to the locations of the two food rewards, finches located a food reward more accurately than on the one-trial tasks, and were also more likely to recall the locations of the different food types. In this foraging paradigm, experience and motivation may have influenced the birds’ performance.     

An assessment of memory awareness in tufted capuchin monkeys (Cebus apella)

Humans, apes, and rhesus monkeys demonstrate memory awareness by collecting information when ignorant and acting immediately when informed. In this study, five capuchin monkeys searched for food after either watching the experimenter bait one of four opaque tubes (seen trials), or not watching (unseen trials). Monkeys with memory awareness should look into the tubes before making a selection only on unseen trials because on seen trials they already know the location of the food. In Experiment 1, one of the five capuchins looked significantly more often on unseen trials. In Experiment 2, we ensured that the monkeys attended to the baiting by interleaving training and test sessions. Three of the five monkeys looked more often on unseen trials. Because monkeys looked more often than not on both trial types, potentially creating a ceiling effect, we increased the effort required to look in Experiment 3, and predicted a larger difference in the probability of looking between seen and unseen trials. None of the five monkeys looked more often on unseen trials. These findings provide equivocal evidence for memory awareness in capuchin monkeys using tests that have yielded clear evidence in humans, apes, and rhesus monkeys.     

Evidence of teaching in atlantic spotted dolphins (Stenella frontalis) by mother dolphins foraging in the presence of their calves

Teaching is a powerful form of social learning, but there is little systematic evidence that it occurs in species other than humans. Using long-term video archives the foraging behaviors by mother Atlantic spotted dolphins (Stenella frontalis) were observed when their calves were present and when their calves were not present, including in the presence of non-calf conspecifics. The nine mothers we observed chased prey significantly longer and made significantly more referential body-orienting movements in the direction of the prey during foraging events when their calves were present than when their calves were not present, regardless of whether they were foraging alone or with another non-calf dolphin. Although further research into the potential consequences for the naïve calves is still warranted, these data based on the maternal foraging behavior are suggestive of teaching as a social-learning mechanism in nonhuman animals.     

How western scrub-jays (Aphelocoma californica) select a nut: effects of the number of options, variation in nut size, and social competition among foragers

Western scrub-jays (Aphelocoma californica) often visually assess and handle several whole (unshelled) peanuts before selecting one to transport and cache; this behavior is a search for a preferred heavy nut. I repeatedly video-taped individually identifiable jays as they landed on a feeding platform and chose from presentations of peanuts that varied in the number of items or in the distribution of sizes. I examined how differences among these presentations and a bird’s social status affected the amount of assessment and the economic consequences of choice. I also examined the specific patterns of handling peanuts, called sampling, to quantify the degree to which sampling sequences were typified by repeated comparisons among sampled peanuts (retrospective sampling), or sequential assessment and rejection of peanuts (prospective sampling). Peanut assessment was more extensive and prospective when there were many options from which to choose than when there were few. Peanut assessment was more extensive and retrospective when options were similar in size than when they varied. Scrub-jays were more likely to make repeated comparisons immediately before selecting a peanut than elsewhere in a sampling sequence. Subordinate scrub-jays, who were at the greatest risk of pre-emption by competitors, assessed peanuts less extensively and were more prospective in their sampling than dominants. Unless peanuts were very similar in size, jays were more accurate at selecting a high-quality peanut and achieved a higher rate of food storage than if they had not assessed. These results show that scrub-jays can adaptively modify how they search to improve their rate of food storage, and also suggest some of the specific search tactics used by jays when assessing peanuts. Received: 26 April 1999 / Accepted after revision: 10 October 1999     

Cuttlefish (Sepia officinalis: Cephalopoda) hunting behavior and associative learning

Because most learning studies in cephalopods have been performed on octopods, it remains unclear whether such abilities are specific to octopus, or whether they correlate with having a larger and more centrally organized brain. To investigate associative learning in a different cephalopod, six sexually mature cuttlefish (Sepia officinalis) participated in a counterbalanced, within-subjects, appetitive, classical conditioning procedure. Two plastic spheres (conditioned stimuli, CSs), differing in brightness, were presented sequentially. Presentation of the CS+ was followed 5 s later by a live feeder fish (unconditioned stimulus, US). Cuttlefish began to attack the CS+ with the same type of food-acquisition seizures used to capture the feeder fish. After seven blocks of training (42 presentations of each CS) the difference in seizure probability between CS+ and CS– trials more than doubled; and was found to be significantly higher in late versus early blocks. These results indicate that cuttlefish exhibit autoshaping under some conditions. The possible ecological significance of this type of learning is briefly discussed.Electronic Supplementary Material Supplementary material is available for this article at