Showing papers in "Animal Cognition in 2009"
TL;DR: It is concluded that gorilla gestural communication is based on a species-typical repertoire, like those of most other mammalian species but very much larger, and employed for intentional communication to specific individuals.
Abstract: Social groups of gorillas were observed in three captive facilities and one African field site. Cases of potential gesture use, totalling 9,540, were filtered by strict criteria for intentionality, giving a corpus of 5,250 instances of intentional gesture use. This indicated a repertoire of 102 gesture types. Most repertoire differences between individuals and sites were explicable as a consequence of environmental affordances and sampling effects: overall gesture frequency was a good predictor of universality of occurrence. Only one gesture was idiosyncratic to a single individual, and was given only to humans. Indications of cultural learning were few, though not absent. Six gestures appeared to be traditions within single social groups, but overall concordance in repertoires was almost as high between as within social groups. No support was found for the ontogenetic ritualization hypothesis as the chief means of acquisition of gestures. Many gestures whose form ruled out such an origin, i.e. gestures derived from species-typical displays, were used as intentionally and almost as flexibly as gestures whose form was consistent with learning by ritualization. When using both classes of gesture, gorillas paid specific attention to the attentional state of their audience. Thus, it would be unwarranted to divide ape gestural repertoires into ‘innate, species-typical, inflexible reactions’ and ‘individually learned, intentional, flexible communication’. We conclude that gorilla gestural communication is based on a species-typical repertoire, like those of most other mammalian species but very much larger. Gorilla gestures are not, however, inflexible signals but are employed for intentional communication to specific individuals.
237 citations
TL;DR: It is observed that 6-month-old infants showed a wider tendency for left gaze preference towards objects and faces of different species and orientation, while in adults the bias appears only towards upright human faces, and face- and species-sensitive gaze asymmetry is more widespread in the animal kingdom than previously recognised.
Abstract: While viewing faces, human adults often demonstrate a natural gaze bias towards the left visual field, that is, the right side of the viewee’s face is often inspected first and for longer periods. Using a preferential looking paradigm, we demonstrate that this bias is neither uniquely human nor limited to primates, and provide evidence to help elucidate its biological function within a broader social cognitive framework. We observed that 6-month-old infants showed a wider tendency for left gaze preference towards objects and faces of different species and orientation, while in adults the bias appears only towards upright human faces. Rhesus monkeys showed a left gaze bias towards upright human and monkey faces, but not towards inverted faces. Domestic dogs, however, only demonstrated a left gaze bias towards human faces, but not towards monkey or dog faces, nor to inanimate object images. Our findings suggest that face- and species-sensitive gaze asymmetry is more widespread in the animal kingdom than previously recognised, is not constrained by attentional or scanning bias, and could be shaped by experience to develop adaptive behavioural significance.
201 citations
TL;DR: It is suggested that 3-year-old children are able to rely on the direction of the index finger, and show the strongest ability to generalize to unfamiliar gestures.
Abstract: We investigated whether dogs and 2-, and 3-year-old human infants living, in some respects, in very similar social environments are able to comprehend various forms of the human pointing gesture. In the first study, we looked at their ability to comprehend different arm pointing gestures (long cross-pointing, forward cross-pointing and elbow cross-pointing) to locate a hidden object. Three-year-olds successfully used all gestures as directional cues, while younger children and dogs could not understand the elbow cross-pointing. Dogs were also unsuccessful with the forward cross-pointing. In the second study, we used unfamiliar pointing gestures i.e. using a leg as indicator (pointing with leg, leg cross-pointing, pointing with knee). All subjects were successful with leg pointing gestures, but only older children were able to comprehend the pointing with knee. We suggest that 3-year-old children are able to rely on the direction of the index finger, and show the strongest ability to generalize to unfamiliar gestures. Although some capacity to generalize is also evident in younger children and dogs, especially the latter appear biased in the use of protruding body parts as directional signals.
137 citations
TL;DR: It is concluded that innovators stimulate others to experiment with the solutions they display, but that chimpanzees are nevertheless conservative; mastery of a skill inhibits further exploration, and hence adoption of alternative techniques even if these are more efficient.
Abstract: Geographic variation in socially transmitted skills and signals, similar to human culture, has been well documented for great apes. The rules governing the adoption of novel behaviours, however, are still largely unknown. We conducted an innovation-and-transmission experiment with two groups of chimpanzees living at hopE Primate Sanctuary Ganserndorf, Austria, presenting a board on which food had to be manoeuvred around obstacles to be acquired. Most chimpanzees used sticks to acquire the food, but five adults independently invented a novel technique, rattling, which was subsequently tested by almost all group members. However, individuals who had become proficient with sticks were reluctant to switch to rattling, despite it being more efficient. Similarly, after rattling was prevented, rattle specialists kept trying to rattle and made no attempt to use the stick technique, despite their knowledge about its existence. We conclude that innovators stimulate others to experiment with the solutions they display, but that chimpanzees are nevertheless conservative; mastery of a skill inhibits further exploration, and hence adoption of alternative techniques even if these are more efficient. Consequently, conformity among group members should not be expected in great apes when individuals develop proficiency at different techniques. Conservatism thus joins conformity as a mechanism to bring about cultural uniformity and stability.
126 citations
TL;DR: It is concluded that contingent reciprocity does not spontaneously arise in experimental settings, despite the fact that patterns of behavior in the field indicate that individuals cooperate preferentially with reciprocating partners.
Abstract: Chimpanzees provide help to unrelated individuals in a broad range of situations. The pattern of helping within pairs suggests that contingent reciprocity may have been an important mechanism in the evolution of altruism in chimpanzees. However, correlational analyses of the cumulative pattern of interactions over time do not demonstrate that helping is contingent upon previous acts of altruism, as required by the theory of reciprocal altruism. Experimental studies provide a controlled approach to examine the importance of contingency in helping interactions. In this study, we evaluated whether chimpanzees would be more likely to provide food to a social partner from their home group if their partner had previously provided food for them. The chimpanzees manipulated a barpull apparatus in which actors could deliver rewards either to themselves and their partners or only to themselves. Our findings indicate that the chimpanzees’ responses were not consistently influenced by the behavior of their partners in previous rounds. Only one of the 11 dyads that we tested demonstrated positive reciprocity. We conclude that contingent reciprocity does not spontaneously arise in experimental settings, despite the fact that patterns of behavior in the field indicate that individuals cooperate preferentially with reciprocating partners.
121 citations
TL;DR: It is shown that magpies can remember which food item they hoarded where, and when, even if the food items only differ from each other in their colour and are dispersed throughout a continuous caching substrate.
Abstract: Some animals have been shown to be able to remember which type of food they hoarded or encountered in which location and how long ago (what-where-when memory). In this study, we test whether magpies (Pica pica) also show evidence of remembering these different aspects of a past episode. Magpies hid red- and blue-dyed pellets of scrambled eggs in a large tray containing wood shavings. They were allowed to make as many caches as they wanted. The birds were then returned either the same day or the next day to retrieve the pellets. If they returned the same day, one colour of pellets was replaced with wooden beads of similar size and colour, while if they returned the next day this would happen to the other colour. Over just a few trials, the birds learned to only search for the food pellets, and ignore the beads, of the appropriate colour for the given retention interval. A probe trial in which all items were removed showed that the birds persisted in searching for the pellets and not the beads. This shows that magpies can remember which food item they hoarded where, and when, even if the food items only differ from each other in their colour and are dispersed throughout a continuous caching substrate.
119 citations
TL;DR: Research of domestic dog behavior in a natural setting shows sensitivity to the visual attention of their partners when engaged in dyadic rough-and-tumble play, which shows attention to, and acted to manipulate, a feature of other dogs that mediates their ability to respond: which feature in human interaction is called “attention”.
Abstract: The social cognitive capacities of dogs, including their communication skills and use of visual attention cues, have recently been investigated in numerous experimental studies. This paper reports on research of domestic dog behavior in a natural setting, which shows sensitivity to the visual attention of their partners when engaged in dyadic rough-and-tumble play. The sequential behaviors and head-direction of both dogs were noted throughout the bouts. The behaviors were differentially used according to the partner’s posture. Play signals were sent nearly exclusively to forward-facing conspecifics; attention-getting behaviors were used most often when a playmate was facing away, and before signaling an interest to play. In addition, the mode of attention-getter matched the degree of inattentiveness of the playmate: stronger attention-getters were used when a playmate was looking away or distracted, less forceful ones when the partner was facing forward or laterally. In other words, these dogs showed attention to, and acted to manipulate, a feature of other dogs that mediates their ability to respond: which feature in human interaction is called “attention”.
109 citations
TL;DR: The results, analyzed at both the group and the individual levels, showed no difference in the performance according to age, indicating that in dogs the comprehension of the human pointing may require only very limited and rapid early learning to fully develop.
Abstract: In spite of the rather different procedures actually used in comparative studies to test the ability of different species to rely on the human pointing gesture, there is no debate on the high performance of dogs in such tasks. Very little is known, however, on the course through which they acquire this ability or the probable factors influencing the process. Important developmental questions have remained unsolved and also some methodological concerns should be addressed before we can convincingly argue for one interpretation or another. In this study we tested 180 dogs of different age (from 2 months to adults) to investigate their performance in the human distal momentary pointing gesture. The results, analyzed at both the group and the individual levels, showed no difference in the performance according to age, indicating that in dogs the comprehension of the human pointing may require only very limited and rapid early learning to fully develop. Interestingly, neither the keeping conditions nor the time spent in active interaction with the owner, and not even some special (agility) training for using human visual cues, had significant effect on the success and explained individual differences. The performance of the dogs was rather stable over time: during the 20 trials within a session and even when subsamples of different age were repeatedly tested. Considering that in spite of the general success at the group level, more than half of the dogs were not successful at the individual level, we revealed alternative “decision-making rules” other than following the pointing gesture of the experimenter.
108 citations
TL;DR: Horses spontaneously selected the greater of the two quantities when the numerosities were small, adding to evidence showing spontaneous quantity assessment in a variety of species.
Abstract: The ability to select the greater numerosity over another in small sets seems to stem from the calculation of which set contains more, and has been taken as evidence of a primordial representation at the roots of the primate numerical system. We tested 56 horses (Equus caballus) in a paradigm previously used with human infants and nonhuman primates. Horses saw two quantities paired in contrasts—2 versus 1, 3 versus 2, 6 versus 4 and a control for volume, 2 versus 1 big—and had to make a choice by snout touching the container holding the numerosity selected. The horses spontaneously selected the greater of the two quantities when the numerosities were small. These results add to evidence showing spontaneous quantity assessment in a variety of species.
103 citations
TL;DR: The first analysis of repeated reuse of the same tool-composites in wild chimpanzees is presented, which may reflect an ability to recognise the nut-cracker as a single tool, as well as discrimination of tool quality-effectiveness.
Abstract: Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuable data towards elucidating the variables that influenced the emergence and development of the first lithic industries among Plio-Pleistocene hominins. Such data help to identify potential behaviours entailed in the first percussive technologies that are invisible in archaeological records. The long-term research site of Bossou in Guinea features a unique chimpanzee community whose members systematically use portable stones as hammers and anvils to crack open nuts in natural as well as in field experimental settings. Here we present the first analysis of repeated reuse of the same tool-composites in wild chimpanzees. Data collected over 5 years of experimental nut-cracking sessions at an “outdoor laboratory” site were assessed for the existence of systematic patterns in the selection of tool-composites, at group and at individual levels. Chimpanzees combined certain stones as hammer and anvil more often than expected by chance, even when taking into account preferences for individual stones by themselves. This may reflect an ability to recognise the nut-cracker as a single tool (composed of two elements, but functional only as a whole), as well as discrimination of tool quality-effectiveness. Through repeatedly combining the same pairs of stones—whether due to preferences for particular composites or for the two elements independently—tool-users may amplify use-wear traces and increase the likelihood of fracturing the stones, and thus of detaching pieces by battering.
100 citations
TL;DR: Results corroborate novel object research conducted in captive populations and suggest that baboons and geladas have differences in behavioral flexibility that have been shaped by ecological (rather than social) differences between the two species.
Abstract: Behavioral flexibility is considered by some to be one of the hallmarks of advanced cognitive ability. One measure of behavioral flexibility is how subjects respond to novel objects. Despite growing interest in comparative cognition, no comparative research on neophilia in wild primates has been conducted. Here, we compare responses to novel objects in wild chacma baboons (Papio ursinus) and geladas (Theropithecus gelada). Baboons and geladas are closely related taxa, yet they differ in their ecology and degree of social tolerance: (1) baboons are habitat and dietary generalists, whereas geladas have one of the most specialized primate diets (90% grass); (2) baboons exhibit an aversion toward extra-group individuals, whereas geladas typically exhibit an attraction toward them. Using subjects of all age and sex classes, we examined responses to three different objects: a plastic doll, a rubber ball, and a metal can. Overall, baboon subjects exhibited stronger responses to the objects (greater neophilia and exploration) than gelada subjects, yet we found no evidence that the geladas were afraid of the objects. Furthermore, baboons interacted with the objects in the same way they might interact with a potential food item. Responses were unrelated to sex, but immatures showed more object exploration than adults. Results corroborate novel object research conducted in captive populations and suggest that baboons and geladas have differences in behavioral flexibility (at least in this cognitive domain) that have been shaped by ecological (rather than social) differences between the two species.
TL;DR: These findings provide equivocal evidence for memory awareness in capuchin monkeys using tests that have yielded clear evidence in humans, apes, and rhesus monkeys.
Abstract: 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.
TL;DR: The important role of mothers and learning opportunity in the acquisition of a hazardous tool-use behaviour is demonstrated and it is suggested that chimpanzee material culture is a product of a complex interaction between social processes and ecological factors.
Abstract: We currently have little understanding of the influence of learning opportunity, whether social or environmental, and maternal role on tool-use acquisition in young wild chimpanzees. This study aims to fill this gap by focusing on the acquisition of ant-dipping among chimpanzees of Bossou, Guinea. Ant-dipping is a hazardous tool-use behaviour aimed at army ants (Dorylus spp.). Bossou chimpanzees target these ants both at nests (high risk) and trails (low risk) and employ two techniques to consume them: direct mouthing and pull-through. We present data for 13 mother-offspring pairs (1-10 years old). Mothers with young < or =5 years old dipped significantly more often at trails than at nests, thus minimizing the risk posed to themselves and their young. Infants thus benefited from better conditions to observe and practice ant-dipping. Mothers also varied greatly in their percent time spent ant-dipping and offspring differed in their learning opportunity. Our results suggest that high opportunity young started to observe and perform ant-dipping sooner and were better at ant-dipping than low opportunity young. Although mothers and weaned offspring correlated positively in their percent time spent dipping and proficiency, they did not match in technique used or tool length. Finally, we propose that the learning trajectory of young may predict individual and sex differences in adulthood. This study demonstrates the important role of mothers and learning opportunity in the acquisition of a hazardous tool-use behaviour and suggests that chimpanzee material culture is a product of a complex interaction between social processes and ecological factors.
TL;DR: The metamemory possessed by this new world monkey species may be more like a flag, showing strength of memory trace, than an elaborate representation showing details of the memory trace.
Abstract: Whereas evidence for metacognition by nonhuman primates has been obtained in great apes and old world monkeys, it is weaker in new world monkeys. For instance, capuchin monkeys may fail to recognize their own knowledge of the location of invisible bait. In the present study, we tested whether tufted capuchin monkeys would flexibly change their behavior in a delayed matching-to-sample (DMTS) test depending upon the strength of their memory trace of the sample. In Experiment 1, two monkeys were tested on a modified 9-alternative DMTS task with various delays on a computerized display. In some trials, the monkeys could choose whether to go for a memory test or for a simple key touch as an escape from the test. In other trials, they were forced to go for the memory test. Both monkeys escaped from the memory test more often when their matching accuracy on forced tests was lower. In one of the monkeys, the matching accuracies on chosen memory tests decreased more slowly as a function of delay length, and were higher after long delays than those on forced memory tests. This suggests that at least one capuchin monkey was able to recognize the strength of his own memory trace. Experiment 2 employed occasional no-sample tests, in which the monkeys faced the task choice without presentation of any sample for the trial. The monkey who was successful in Experiment 1 declined the memory test more often in no-sample trials than regular trials, further indicating metamemory in this individual. In Experiment 3, this successful monkey received a task, in which he was sometimes able to choose between shape MTS or texture MTS tasks. However, his matching accuracies did not differ between chosen tasks and forced tasks. Thus, the metamemory possessed by this new world monkey species may be more like a flag, showing strength of memory trace, than an elaborate representation showing details of the memory trace.
TL;DR: This study shows that forest chimpanzees possess a precise spatial memory which allows them to remember the location of numerous resources and use this information to select the most attractive resources.
Abstract: It is assumed that spatial memory contributes crucially to animal cognition since animals’ habitats entail a large number of dispersed and unpredictable food sources. Spatial memory has been investigated under controlled conditions, with different species showing and different conditions leading to varying performance levels. However, the number of food sources investigated is very low compared to what exists under natural conditions, where food resources are so abundant that it is difficult to precisely identify what is available. By using a detailed botanical map containing over 12,499 trees known to be used by the Tai chimpanzees, we created virtual maps of all productive fruit trees to simulate potential strategies used by wild chimpanzees to reach resources without spatial memory. First, we simulated different assumptions concerning the chimpanzees’ preference for a particular tree species, and, second, we varied the detection field to control for the possible use of smell to detect fruiting trees. For all these assumptions, we compared simulated distance travelled, frequencies of trees visited, and revisit rates with what we actually observed in wild chimpanzees. Our results show that chimpanzees visit rare tree species more frequently, travel shorter distances to reach them, and revisit the same trees more often than if they had no spatial memory. In addition, we demonstrate that chimpanzees travel longer distances to reach resources where they will eat for longer periods of time, and revisit resources more frequently where they ate for a long period of time during their first visit. Therefore, this study shows that forest chimpanzees possess a precise spatial memory which allows them to remember the location of numerous resources and use this information to select the most attractive resources.
TL;DR: The results show that memory for what, where, and when occurs in food-storing birds outside the corvid family, does not require food caching or retrieval, and that remembering “when” can depend on the nature of the task.
Abstract: Episodic memory is the ability to remember personally experienced past events (Tulving in Organization of memory. Academic Press, San Diego, pp. 381–403, 1972). In non-human animals, the behavioural criterion for episodic-like memory is remembering “what” occurred in conjunction with “when” and “where” (Clayton and Dickinson in Nature 395:272–274, 1998). We conducted tests for “what, where, and when” memory in a food-storing bird, the black-capped chickadee (Poecile atricapillus). In Experiment 1, chickadees found sunflower seeds and mealworms in concealed sites in their home cage. Birds later re-visited these sites after either a short (3 h) or long (123 h) retention interval. Chickadees normally prefer mealworms, but at the long retention interval mealworms were degraded in taste and appearance. Chickadees showed some memory for what kind of food they had encountered and where, but no memory for when food had previously been found. Experiment 2 followed a similar procedure, except that chickadees searched for hidden sunflower seeds and mealworms in trees in an indoor aviary. These more natural conditions increased both the spatial scale of the task and the effort required to find food. In this experiment, birds showed evidence for all three components of what–where–when memory. Unlike some previous studies of episodic-like memory, birds’ behaviour cannot be accounted for by differential memory strength for more recent events. The results show that memory for what, where, and when occurs in food-storing birds outside the corvid family, does not require food caching or retrieval, and that remembering “when” can depend on the nature of the task.
TL;DR: Data based on the maternal foraging behavior of mother Atlantic spotted dolphins are suggestive of teaching as a social-learning mechanism in nonhuman animals.
Abstract: 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 naive calves is still warranted, these data based on the maternal foraging behavior are suggestive of teaching as a social-learning mechanism in nonhuman animals.
TL;DR: These results are, to the authors' knowledge, the first to demonstrate auditory laterality in an ungulate species and open further lines of thought on the influence of the social “value” of calls and the listener’s arousal on auditory processing and laterality.
Abstract: Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener’s arousal on auditory processing and laterality.
TL;DR: It is found chimpanzees looked longer at severe victim screams than either mild victim screams or tantrum screams, indicating that chimpanzees attend to the informational content of screams and are able to distinguish between different scream variants, which form part of a graded continuum.
Abstract: When experiencing aggression from group members, chimpanzees commonly produce screams. These agonistic screams are graded signals and vary acoustically as a function of the severity of aggression the caller is facing. We conducted a series of field playback experiments with a community of wild chimpanzees in the Budongo Forest, Uganda, to determine whether individuals could meaningfully distinguish between screams given in different agonistic contexts. We compared six subjects’ responses to screams given in response to severe and mild aggression. Subjects consistently discriminated between the two scream types. To address the possibility that the response differences were driven directly by the screams’ peripheral acoustic features, rather than any attached social meaning, we also tested the subjects’ responses to tantrum screams. These screams are given by individuals that experienced social frustration, but no physical threat, yet acoustically they are very similar to screams of victims of severe aggression. We found chimpanzees looked longer at severe victim screams than either mild victim screams or tantrum screams. Our results indicate that chimpanzees attend to the informational content of screams and are able to distinguish between different scream variants, which form part of a graded continuum.
TL;DR: Examination of acquisition of techniques used to process Luehea candida fruits in a wild population of white-faced capuchin monkeys, Cebus capucinus, residing in and near Lomas Barbudal Biological Reserve, Costa Rica revealed that the technique most frequently observed significantly predicted the technique adopted by female observers, particularly in the second year of life.
Abstract: Researchers of “culture” have long been interested in the role of social learning in establishing patterns of behavioral variation in wild animals, but very few studies examine this issue using a developmental approach. This 7-year study examines the acquisition of techniques used to process Luehea candida fruits in a wild population of white-faced capuchin monkeys, Cebus capucinus, residing in and near Lomas Barbudal Biological Reserve, Costa Rica. The two techniques for extracting seeds (pounding or scrubbing) were approximately equal in efficiency, and subjects experimented with both techniques before settling on one technique—typically the one they most frequently observed. In a sample of 106 subjects that had already settled on a preferred technique, the females adopted the maternal technique significantly more often than expected by chance, but the males did not. Using a longitudinal approach, I examined the acquisition of Luehea processing techniques during the first 5 years of life. Regression analysis revealed that the technique most frequently observed (measured as proportion of Luehea processing bouts observed that used pounding as opposed to scrubbing) significantly predicted the technique adopted by female observers, particularly in the second year of life; the amount of impact of the observed technique on the practiced technique was somewhat less significant for male observers. These results held true for (a) observations of maternal technique only, (b) observations of technique used by all individuals other than the mother, and (c) observations of maternal and non-maternal techniques combined.
TL;DR: The potential existence of individual acoustic signatures in whinny calls and the ability of horses to discriminate by ear individuals varying in their degree of familiarity are investigated and a process of vocal coding/decoding of information is suggested.
Abstract: Many animal species that rely mainly on calls to communicate produce individual acoustic structures, but we wondered whether individuals of species better known as visual communicants, with small vocal repertoires, would also exhibit individual distinctiveness in calls. Moreover, theoretical advances concerning the evolution of social intelligence are usually based on primate species data, but relatively little is known about the social cognitive capacities of non-primate mammals. However, some non-primate species demonstrate auditory recognition of social categories and possess mental representation of their social network. Horses (Equus caballus) form stable social networks and although they display a large range of visual signals, they also use long-distance whinny calls to maintain contact. Here, we investigated the potential existence of individual acoustic signatures in whinny calls and the ability of horses to discriminate by ear individuals varying in their degree of familiarity. Our analysis of the acoustic structure of whinnies of 30 adult domestic horses (ten stallions, ten geldings, ten mares) revealed that some of the frequency and temporal parameters carried reliable information about the caller’s sex, body size and identity. However, no correlations with age were found. Playback experiments evaluated the behavioural significance of this variability. Twelve horses heard either control white noise or whinnies emitted by group members, familiar neighbours or unfamiliar horses. While control sounds did not induce any particular response, horses discriminated the social category of the callers and reacted with a sound-specific behaviour (vigilance and attraction varied with familiarity). Our results support the existence of social knowledge in horses and suggest a process of vocal coding/decoding of information.
TL;DR: The hypothesis that horses can form a memory of humans that impacts their reactions in subsequent interactions supports the hypothesis that perception of humans by horses may be based on experience, i.e. repeated interactions.
Abstract: This study aimed to determine whether horses have a kind of memory of humans (based on previous interactions), leading to a general significance of humans revealed by their reactions to humans in subsequent interactions. Subjects were 59 adult horses used to interact daily with humans. Three types of behavioural tests involving an unknown experimenter evaluated three possibly different memorized types of human–animal interactions (not work-related, using work-related objects, unfamiliar working task). We also performed standardized observations of routine interactions between each horse and its familiar handler (caretaker). To get a broad overview of the horses’ reactions to humans, we recorded both investigative and aggressive behaviours during the tests, representing respectively a “positive” and a “negative” memory of the relationship. Whereas correlations between tests revealed a general perception of humans as either positive or negative, unusual tests, i.e. that are not usually performed, elicited more positive reactions. Moreover, some horses reacted positively to a motionless person in their box, but negatively when this same person approached them, for example for halter fitting. Overall, aggressive reactions were more reliable indicators of the relationship than positive reactions, both between tests and between familiar and unfamiliar humans. Our results also show generalization of the perception of humans. These results support our hypothesis that perception of humans by horses may be based on experience, i.e. repeated interactions. Altogether, our results support the hypothesis that horses can form a memory of humans that impacts their reactions in subsequent interactions.
TL;DR: Monkeys use both second-order configural and feature-based processing to recognize the faces of conspecifics, but they use primarily feature- based strategies to recognize human faces.
Abstract: Successful integration of individuals in macaque societies suggests that monkeys use fast and efficient perceptual mechanisms to discriminate between conspecifics. Humans and great apes use primarily holistic and configural, but also feature-based, processing for face recognition. The relative contribution of these processes to face recognition in monkeys is not known. We measured face recognition in three monkeys performing a visual paired comparison task. Monkey and humans faces were (1) axially rotated, (2) inverted, (3) high-pass filtered, and (4) low-pass filtered to isolate different face processing strategies. The amount of time spent looking at the eyes, mouth, and other facial features was compared across monkey and human faces for each type of stimulus manipulation. For all monkeys, face recognition, expressed as novelty preference, was intact for monkey faces that were axially rotated or spatially filtered and was supported in general by preferential looking at the eyes, but was impaired for inverted faces in two of the three monkeys. Axially rotated, upright human faces with a full range of spatial frequencies were also recognized, however, the distribution of time spent exploring each facial feature was significantly different compared to monkey faces. No novelty preference, and hence no inferred recognition, was observed for inverted or low-pass filtered human faces. High-pass filtered human faces were recognized, however, the looking pattern on facial features deviated from the pattern observed for monkey faces. Taken together these results indicate large differences in recognition success and in perceptual strategies used by monkeys to recognize humans versus conspecifics. Monkeys use both second-order configural and feature-based processing to recognize the faces of conspecifics, but they use primarily feature-based strategies to recognize human faces.
TL;DR: Interestingly, elephants did not exhibit disparity or magnitude effects, in which performance declines with a smaller difference between quantities in a two-choice task, or the total quantity increases, respectively, suggesting that elephants may be using a different mechanism to compare and represent quantities than previously suggested for other species.
Abstract: This study investigated whether Asian elephants can make relative quantity judgment (RQJ), a dichotomous judgment of unequal quantities ordered in magnitude. In Experiment 1, elephants were simultaneously shown two baskets with differing quantities of bait (up to 6 items). In Experiment 2, elephants were sequentially presented with baits, which could not be seen by elephants in their total quantities. The task of elephants was to choose the larger quantity in both experiments. Results showed that the elephants chose the larger quantity with significantly greater frequency. Interestingly, the elephants did not exhibit disparity or magnitude effects, in which performance declines with a smaller difference between quantities in a two-choice task, or the total quantity increases, respectively. These findings appear to be inconsistent with the previous reports of RQJ in other animals, suggesting that elephants may be using a different mechanism to compare and represent quantities than previously suggested for other species.
TL;DR: These results unite known patterns of risk-sensitive decision making in human and nonhuman primates and suggest that factors such as the way a decision is framed or internal state may underlie observed variation in risk preferences between and within species.
Abstract: Determining how both humans and animals make decisions in risky situations is a central problem in economics, experimental psychology, behavioral economics, and neurobiology. Typically, humans are risk seeking for gains and risk averse for losses, while animals may display a variety of preferences under risk depending on, amongst other factors, internal state. Such differences in behavior may reflect major cognitive and cultural differences or they may reflect differences in the way risk sensitivity is probed in humans and animals. Notably, in most studies humans make one or a few choices amongst hypothetical or real monetary options, while animals make dozens of repeated choices amongst options offering primary rewards like food or drink. To address this issue, we probed risk-sensitive decision making in human participants using a paradigm modeled on animal studies, in which rewards were either small squirts of Gatorade or small amounts of real money. Possible outcomes and their probabilities were not made explicit in either case. We found that individual patterns of decision making were strikingly similar for both juice and for money, both in overall risk preferences and in trial-to-trial effects of reward outcome on choice. Comparison with decisions made by monkeys for juice in a similar task revealed highly similar gambling styles. These results unite known patterns of risk-sensitive decision making in human and nonhuman primates and suggest that factors such as the way a decision is framed or internal state may underlie observed variation in risk preferences between and within species.
TL;DR: Failure- and latency-to-exchange measures showed that witnesses were unaffected by the food type offered to models (no inequity aversion) and models were unaffectedBy their history of food type offer (no frustration).
Abstract: Brosnan and de Waal (Nature 425:297-299, 2003) reported that if a witness monkey saw a model monkey receive a high-value food, the witness was more inclined to reject a previously acceptable, but low-value food. Later work demonstrated that this alleged inequity aversion might be due to frustration induced by switching subjects from their role as models receiving a high-value food to the role of witnesses receiving a low-value food. In the present study, pairs of female capuchins exchanged a token for either a high- or a low-value food without switching their model-witness roles. Witnesses could exchange a token for a low-value food after an adjacent model had exchanged a token for the same food (Equity Condition) or for a high-value food (Inequity Condition). Failure- and latency-to-exchange measures showed that witnesses were unaffected by the food type offered to models (no inequity aversion). Moreover, models were unaffected by their history of food type offered (no frustration). These results join earlier work suggesting that alleged inequity effects depend on frustration-induction procedures. Furthermore, inequity effects sometimes fail to emerge because frustration induction in nonhuman primates is labile.
TL;DR: These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time.
Abstract: Three horses (Equus caballus) with a history of performing cognitive tasks including discrimination learning, categorization, and concept use were tested to evaluate their long-term memory (LTM) in three experiments. In addition, use of LCD multi-displays for stimulus presentation was incorporated into cognition testing protocol for the first time with horses. Experiment 1 tested LTM for discrimination learning that originally occurred 6 years earlier. Five sets of stimuli were used and the two horses tested showed no decrement in performance on four of the sets; however, both horses did score below chance on one set. Experiment 2 examined long-term categorization recall 10 years after horses had demonstrated the ability to make stimulus selections based on shared characteristics within a given category. The horse tested for LTM after the decade-long interval immediately and consistently applied the previously learned categorization rule to not only familiar but also novel sets of stimuli. Experiment 3 tested another horse for LTM for a relative size concept. This horse had originally demonstrated concept rule use in order to select stimuli based on their relative size to one another. More than 7 years later and without further training, this horse reliably applied the previously established size concept to both familiar and novel sets of stimuli. These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time. These studies also demonstrate the adaptive nature of horses with regard to their ability to generalize over several different testing conditions.
TL;DR: The percentage of dogs showing contagious yawning was less than chimpanzees and humans showing this behavior, and considerably less than a recently published report investigating this behavior in dogs.
Abstract: We report an experimental investigation into whether domesticated dogs display contagious yawning. Fifteen dogs were shown video clips of (1) humans and (2) dogs displaying yawns and open-mouth expressions (not yawns) to investigate whether dogs showed contagious yawning to either of these social stimuli. Only one dog performed significantly more yawns during or shortly after viewing yawning videos than to the open-mouth videos, and most of these yawns occurred to the human videos. No dogs showed significantly more yawning to the open-mouth videos (human or dog). The percentage of dogs showing contagious yawning was less than chimpanzees and humans showing this behavior, and considerably less than a recently published report investigating this behavior in dogs (Joly-Mascheroni et al. in Biol Lett 4:446-448, 2008).
TL;DR: It is shown that lemurs’ gaze following behaviour and ultimate choice are closely linked, and a candidate system for the evolutionary origins of more complex gaze following is proposed: ‘gaze priming.’
Abstract: Although primates have often been found to co-orient visually with other individuals, members of these same species have usually failed to use co-orientation to find hidden food in object-choice experiments. This presents an evolutionary puzzle: what is the function of co-orientation if it is not used for a function as basic as locating resources? Co-orientation responses have not been systematically investigated in object-choice experiments, and requiring co-orientation with humans (as is typical in object-choice tasks) may underestimate other species’ abilities. Using an object-choice task with conspecific models depicted in photographs, we provide experimental evidence that two lemur species (Eulemur fulvus, n = 4, and Eulemur macaco, n = 2) co-orient with conspecifics. Secondly, by analysing together two measures that have traditionally been examined separately, we show that lemurs’ gaze following behaviour and ultimate choice are closely linked. Individuals were more likely to choose correctly after having looked in the same direction as the model, and thus chose objects correctly more often than chance. We propose a candidate system for the evolutionary origins of more complex gaze following: ‘gaze priming.’
TL;DR: It is shown that males can discriminate sources of odours reflecting 1 versus 4 and 1 versus 3 females, but not 2 versus 4 or 1 versus 2, indicating that T. molitor males exhibit a marked preference for sources reflecting more female donors only when numerosity ratios are below 1:2.
Abstract: Numerosity discrimination, the ability to distinguish between sets with more and less items, is recognised as the foundation for higher numerical abilities. Understanding numerosity discrimination from a comparative perspective is hence pivotal in tracing the evolution of numerical representation systems. However, numerosity discrimination has been well studied only in vertebrates, where two innate systems of number representation have been described: an ‘analog magnitude system’ used to discriminate among numerosities by representing them as cardinal magnitudes and a ‘parallel individualisation system’ that allows precise discrimination among small arrays of items (≤4) by representing objects individually. We investigated the existence of quantity discrimination in an insect species (Tenebrio molitor) by using a spontaneous two-choice procedure in which males were exposed to substrates bearing odours from different numbers of females (≤4) in increasing numerosity ratios (1:4, 1:3 and 1:2). We show that males can discriminate sources of odours reflecting 1 versus 4 and 1 versus 3 females, but not 2 versus 4 or 1 versus 2, indicating that T. molitor males exhibit a marked preference for sources reflecting more female donors only when numerosity ratios are below 1:2. We discuss the functional significance of this finding and whether our pattern of results could be best explained by summation of a non-numerical continuous variable or by the existence of a numerosity discrimination mechanism with an operational signature ratio of 1:2.