String‐pulling behaviour in a Harris's Hawk Parabuteo unicinctus

Variations on the string‐pull experiment have been presented to a variety of avian species. Here, we present the results of a basic vertical string‐pull task with a Harris's Hawk Parabuteo unicinctus. A 2‐year‐old subject retrieved a shielded food reward within 8 min on each of eight trials and spontaneously used solving techniques similar to corvids and parrots. Our data contribute to the small body of literature on raptor cognition by showing that it may be within the realm of at least one bird of prey species to perform the string‐pull task similarly to avian species renowned for their high cognitive abilities.     

Testing Problem Solving in Ravens: String-Pulling to Reach Food

The aim of our study was to re‐examine the acquisition of problem‐solving behaviour in ravens: accessing meat suspended from a perch by a string. In contrast to a previous study, here we: (i) controlled for possible effects of fear of the string, competition by dominants, and social learning and (ii) devised a mechanically equivalent but non‐intuitive task to test for the possibility of means–end understanding. One‐year‐old ravens confronted with meat on a string for the first time tried several ways to reach the food. However, five of six birds suddenly performed a coherent sequence of pulling up and stepping on loops of string, essential for solving the problem. Those five birds were also successful in the non‐intuitive task where they had to pull down the string to lift the meat. A second group of birds with similar exposure to strings but without any experience in pulling up meat failed the pull‐down test. These results support the idea that the ravens’ behaviour in accessing meat on a string is not only a product of rapid learning but may involve some understanding of cause–effect relation between string, food and certain body parts.     

Paying attention pays off: Kea improve in loose-string cooperation by attending to partner

Previous studies showed that kea are able to cooperate in experiments based on the loose-string paradigm, but success rates were low, except when tested in stable dyads. We trained kea with low success rates to attend to the handling of the string by a human partner. This vastly improved subsequent coordination during cooperation with kea partners. Furthermore, we tested the kea dyads with a delayed entry set-up (delays of 2, 4 or 6 s) and with two different lengths of string. Increasing length of delay and reducing string length had negative effects on the success rate. This suggests that a complete understanding of the actions was not present as the kea were not able to fully overcome factors that inhibit coordination. Lastly, we tested the three subjects showing the greatest ability to wait for the partner in a partner-choice paradigm: one subject in the central position could pull a loose string with one partner after another in the order it preferred. As a result, the second subjects had much longer waiting times than previously experienced. We could show that the central kea learned to visit both set-ups and that the second partner was able to wait for the partner to arrive before attempting to pull on the string, averaging twice as long as in the standard delay. Taken together, the results from this study highlight that cooperation in the loose-string paradigm is largely dependent on coordination of the subjects. While attention to the partner's actions greatly improved performance, the greater waiting times achieved in the final set-up also suggest that utilizing more ecologically relevant (subject is busy VS subject is held back) delays could further improve the performance of non-human subjects in the delayed loose-string paradigm.     

Cooperative problem solving in a cooperatively breeding primate (Saguinus oedipus)

We investigated cooperative problem solving in unrelated pairs of the cooperatively breeding cottontop tamarin, Saguinus oedipus, to assess the cognitive basis of cooperative behaviour in this species and to compare abilities with other apes and monkeys. A transparent apparatus was used that required extension of two handles at opposite ends of the apparatus for access to rewards. Resistance was applied to both handles so that two tamarins had to act simultaneously in order to receive rewards. In contrast to several previous studies of cooperation, both tamarins received rewards as a result of simultaneous pulling. The results from two experiments indicated that the cottontop tamarins (1) had a much higher success rate and efficiency of pulling than many of the other species previously studied, (2) adjusted pulling behaviour to the presence or absence of a partner, and (3) spontaneously developed sustained pulling techniques to solve the task. These findings suggest that cottontop tamarins understand the role of the partner in this cooperative task, a cognitive ability widely ascribed only to great apes. The cooperative social system of tamarins, the intuitive design of the apparatus, and the provision of rewards to both participants may explain the performance of the tamarins.     

Patterned-String Tasks: Relation between Fine Motor Skills and Visual-Spatial Abilities in Parrots

String-pulling and patterned-string tasks are often used to analyse perceptual and cognitive abilities in animals. In addition, the paradigm can be used to test the interrelation between visual-spatial and motor performance. Two Australian parrot species, the galah (Eolophus roseicapilla) and the cockatiel (Nymphicus hollandicus), forage on the ground, but only the galah uses its feet to manipulate food. I used a set of string pulling and patterned-string tasks to test whether usage of the feet during foraging is a prerequisite for solving the vertical string pulling problem. Indeed, the two species used techniques that clearly differed in the extent of beak-foot coordination but did not differ in terms of their success in solving the string pulling task. However, when the visual-spatial skills of the subjects were tested, the galahs outperformed the cockatiels. This supports the hypothesis that the fine motor skills needed for advanced beak-foot coordination may be interrelated with certain visual-spatial abilities needed for solving patterned-string tasks. This pattern was also found within each of the two species on the individual level: higher motor abilities positively correlated with performance in patterned-string tasks. This is the first evidence of an interrelation between visual-spatial and motor abilities in non-mammalian animals.     

Individuality in Problem Solving: String Pulling in Two Carduelis Species (Aves: Passeriformes)

The research reported here was designed to study the individual peculiarities of birds in solving a problem. Goldfinches Carduelis carduelis and siskins C. spinus were tested with the string‐pulling task: sitting on a perch from which a small food container is suspended by a string the test bird had to lift the container, using the bill to pull the string stepwise up and a foot to hold it, and repeat that until they could reach the food. Fifty‐two goldfinches and 29 siskins raised under controlled conditions were tested individually. Three groups became apparent: ‘inventors’ (23% of goldfinches; 62% of siskins) solved the problem by themselves; ‘imitators’ (25% of goldfinches; 10% of siskins) succeeded after seeing a performing conspecific; ‘duffers’ (52% of goldfinches, 28% of siskins) did not succeed either way. The species – but not the sexes – differed significantly in string‐pulling ability. The results of our experiments indicate that string pulling is an acquired combination of innate behaviour elements. An individual's string‐pulling competence may depend on prior experience of handling branchlets, on trial‐and‐error learning and on social learning (emulation). However, some individuals succeeded without these facilitating factors, while others did not succeed at all despite all of them present. Although functionally and motivationally related to feeding, the learned string pulling is often shown as a playful activity without an obvious reward.     

Parrot behavior at a Rio Manu (Peru) clay lick: temporal patterns,associations, and antipredator responses

Although eating clay at licks (a form of geophagy) has been described, there are few behavioral data on temporal patterns, social interactions, species associations, or reactions to potential predators. We examined the behavior of nine species of macaws, parrots, and parakeets at the Machiguenga Ccolpa, a clay lick on the Rio Manu, Peru in the dry season. Three distinct mixed-species groups used the licks: in the early morning (parrots and small macaws), in mid-morning (large macaws), and in the early afternoon (parakeets), although the latter two groups used the licks at other times of day as well. The first parrots to begin eating at the lick in the early morning were yellow-crowned parrots (Amazona ochrocephala) and dusky-headed parakeets (Aratinga weddellii), followed by blue-headed parrots Pionus sordidus, and then by mealy (Amazona farinosa) and orange-cheeked (Pionopsitta barrabandi) parrots, and chestnut-fronted macaws (Ara severa). Although blue-headed parrots fed in dense groups of over 50, the others rarely exceeded 20 individuals. Scarlet macaws (A. macao) sometimes fed alone or joined the early morning groups, but most associated with a large group of red and green macaws (A. chloroptera) that arrived, often scaring off the smaller birds. On average, about 100 macaws and parrots fed in the early morning, macaw feeding groups averaging just over 40, and the parakeets averaged over 70. Average time at the lick ranged from 28 min for yellow-crowned parrots to 47 min for tui parakeets. Of the early morning group, blue-headed and mealy parrots were the most aggressive and orange-cheeked parrots were the least aggressive. Red and green macaws were more aggressive than scarlet macaws; the parakeets were equally aggressive. All species had more aggressive interactions with conspecifics than with other species. Responses to intruders and predators varied by species of parrot/macaw and type of intruder. In response to intruders or loud calls, responses could be partial (some individuals flew away, circled, and returned), temporary (all individuals flew away but returned within a few minutes), or total (all flew away and abandoned feeding for at least a half hour). The large macaws showed the lowest rate of total abandonment and the parakeets showed the highest. People passing up or down river in boats scared birds from the lick. The local residents (Machiguenga tribespeople in boats) elicited a much greater response than did the researchers. In the recent past, macaws and parrots were hunted for food, feathers, and the pet trade, and the birds response, as well as the presence of parrot and macaw feathers in local villages we visited, suggests some continued exploitation, or a long-term memory in the birds.Communicated by R.F. Oliveira     

Chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) learn to act with other individuals in a cooperative task

We presented two chimpanzees with a task in which they were required to pull each end of a rope simultaneously to drag blocks supporting food into reach. The chimpanzees did not succeed in initial tests. They did not immediately understand the necessity for cooperation, and they did not adjust their behavior to work with the partner. However, the frequency of success gradually increased as the number of sessions increased and the task was varied. They began to look at the partner frequently, wait if the partner was not holding the rope, and pull the rope in synchrony with the partner. However, they did not use interactive behaviors or eye contact to synchronize their behavior. One chimpanzee was then paired with a human partner in the same situation. After initial failures, the chimpanzee began to solicit the human partner for cooperation: looking up at his face, vocalizing, and taking the partner’s hand. When this chimpanzee was again paired with the chimpanzee partner, no soliciting behavior was observed. Thus, the chimpanzees could learn to coordinate their behavior through trial and error. Communicative behavior emerged during the task, but the communication differed according to the identity of the partner.     

Gaze direction - a cue for hidden food in rooks (Corvus frugilegus)?

Other individual's head- and eye-directions can be used as social cues indicating the presence of important events. Among birds, ravens and rooks have been shown to co-orient with conspecifics and with humans by following their gaze direction into distant space and behind visual screens. Both species use screens to cache food in private; also, it had been suggested that they may rely on gaze cues to detect hidden food. However, in an object-choice task, ravens failed to do so, and their competitive lifestyle may have prevented them from relying on these cues. Here we tested closely related and cooperative rooks. Food was hidden in one of two cups and the experimenter gazed at the baited cup. In a second experiment, we aimed to increase the birds’ motivation to choose correctly by increasing the investment needed to obtain the reward. To do so, the birds had to pull on a string to obtain the cup. Here, the birds as a group tended to rely on gaze cues. In addition, individual birds quickly learned to use the cue in both experiments. Although rooks may not use gaze cues to find hidden food spontaneously, they may quickly learn to do so.     

Effects of hunting on guianan forest game birds

To assess the effect of on-going and of previous or near by hunting pressure on game birds in a neotropical rain forest (French Guiana), I compared species abundances between six hunted and disturbed areas, seven non-hunted and pristine areas and eight intermediate areas, undisturbed and not currently hunted, but formerly hunted or close to current hunting areas. I recorded all birds detected within 100 m-wide strip transects, walked at random through every forest type all day long. The frequency of records per 10 h and flock sizes on each transect were averaged over all surveys in each study site (N = 3025 h). Censused groups included terrestrial (tinamous, woodquails, curassow and trumpeter) and arboreal species (guans, macaws, parrots, toucans), but not waterbirds. The abundance of all game birds was higher in non-hunted than in hunted areas, though not always significantly for secretive understorey species (tinamous, guans, wood-quail) or canopy frugivores (parrots, macaws, toucans). The Black Curassow and Gray-winged Trumpeter exhibited the highest and most consistent increases (7–10-fold) and more so between intermediate and non-hunted sites than between hunted and intermediate sites. The mean flock size also increased along the hunting gradient, especially in flocking species (macaws, parrots, trumpeter). Low reproductive rates and/or seasonal movements may explain that hunting pressure could still be felt after hunting has ceased or when it persists only away from a given area. Such a persistent effect would affect some populations in small protected areas.     

What You See Is What You Get? Exclusion Performances in Ravens and Keas

Background

Among birds, corvids and parrots are prime candidates for advanced cognitive abilities. Still, hardly anything is known about cognitive similarities and dissimilarities between them. Recently, exclusion has gained increasing interest in comparative cognition. To select the correct option in an exclusion task, one option has to be rejected (or excluded) and the correct option may be inferred, which raises the possibility that causal understanding is involved. However, little is yet known about its evolutionary history, as only few species, and mainly mammals, have been studied.

Methodology/Principal Findings

We tested ravens and keas in a choice task requiring the search for food in two differently shaped tubes. We provided the birds with partial information about the content of one of the two tubes and asked whether they could use this information to infer the location of the hidden food and adjust their searching behaviour accordingly. Additionally, this setup allowed us to investigate whether the birds would appreciate the impact of the shape of the tubes on the visibility of food. The keas chose the baited tube more often than the ravens. However, the ravens applied the more efficient strategy, choosing by exclusion more frequently than the keas. An additional experiment confirmed this, indicating that ravens and keas either differ in their cognitive skills or that they apply them differently.

Conclusion

To our knowledge, this is the first study to demonstrate that corvids and parrots may perform differently in cognitive tasks, highlighting the potential impact of different selection pressures on the cognitive evolution of these large-brained birds.     

Social factors determine cooperation in marmosets

We investigated which factors determine cooperative behaviour of common marmosets, Callithrix jacchus, in an instrumental task. During the approach phase each of eight individuals of a family group learned in isolation to pull a handle moving a bowl with attractive food towards its reach. In the following two experimental phases we used 16 dyads. In a dyadic training phase we assessed whether the partners were willing to manipulate the apparatus and to share food. In the subsequent cooperation test we examined whether they were willing to cooperate at a slightly modified apparatus whose solution required one individual (the producer) to pull the handle and the other (the scrounger) to grasp the bowl. Although all individuals were willing to cooperate with at least one partner, only half of the dyads solved the task in the cooperation phase. Examination of the factors that correlated with success in this phase revealed that primarily those dyads cooperated in which the dominant subject took the role of the scrounger and the subordinate took the role of the producer. However, in these successful dyads the dominant animal did not force the subordinate partner to pull the handle. Rather, the partners of cooperative dyads shared the reward and pulled equally often in both the dyadic training and the cooperation test. Thus, cooperation of marmosets in an instrumental task seems to depend on a specific distribution of roles and the tolerance of higher-ranking individuals.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Grey parrots use inferential reasoning based on acoustic cues alone

Our ability to make logical inferences is considered as one of the cornerstones of human intelligence, fuelling investigations of reasoning abilities in non-human animals. Yet, the evidence to date is equivocal, with apes as the prime candidates to possess these skills. For instance, in a two-choice task, apes can identify the location of hidden food if it is indicated by a rattling noise caused by the shaking of a baited container. More importantly, they also use the absence of noise during the shaking of the empty container to infer that this container is not baited. However, since the inaugural report of apes solving this task, to the best of our knowledge, no comparable evidence could be found in any other tested species such as monkeys and dogs. Here, we report the first successful and instantaneous solution of the shaking task through logical inference by a non-ape species, the African grey parrot. Surprisingly, the performance of the birds was sensitive to the shaking movement: they were successful with containers shaken horizontally, but not with vertical shaking resembling parrot head-bobbing. Thus, grey parrots seem to possess ape-like cross-modal reasoning skills, but their reliance on these abilities is influenced by low-level interferences.     

Cooperative problem solving in rooks (Corvus frugilegus)

Recent work has shown that captive rooks, like chimpanzees and other primates, develop cooperative alliances with their conspecifics. Furthermore, the pressures hypothesized to have favoured social intelligence in primates also apply to corvids. We tested cooperative problem-solving in rooks to compare their performance and cognition with primates. Without training, eight rooks quickly solved a problem in which two individuals had to pull both ends of a string simultaneously in order to pull in a food platform. Similar to chimpanzees and capuchin monkeys, performance was better when within-dyad tolerance levels were higher. In contrast to chimpanzees, rooks did not delay acting on the apparatus while their partner gained access to the test room. Furthermore, given a choice between an apparatus that could be operated individually over one that required the action of two individuals, four out of six individuals showed no preference. These results may indicate that cooperation in chimpanzees is underpinned by more complex cognitive processes than that in rooks. Such a difference may arise from the fact that while both chimpanzees and rooks form cooperative alliances, chimpanzees, but not rooks, live in a variable social network made up of competitive and cooperative relationships.     

Capuchins do cooperate: the advantage of an intuitive task

We used a cooperative pulling task to examine proximate aspects of cooperation in captive brown capuchin monkeys, Cebus apella. Specifically, our goal was to determine whether capuchins can learn the contingency between their partner's participation in a task and its successful completion. We examined whether the monkeys visually monitored their partners and adjusted pulling behaviour according to their partner's presence. Results on five same-sex pairs of adults indicate that (1) elimination of visual contact between partners significantly decreased success, (2) subjects glanced at their partners significantly more in cooperative tests than in control tests in which no partner-assistance was needed, and (3) they pulled at significantly higher rates when their partner was present rather than absent. Therefore, in contrast to a previous report by Chalmeau et al. (1997, Animal Behaviour, 54, 1215-1225), cooperating capuchins do seem able to take the role of their partner into account. However, the type of task used may be an important factor affecting the level of coordination achieved. Copyright 2000 The Association for the Study of Animal Behaviour.     

DNA sequence analysis to guide the release of blue-and-yellow macaws (Ara ararauna, Psittaciformes, Aves) from the illegal trade back into the wild

The illegal wildlife trade is one of the major threats to Brazil’s biodiversity. Approximately 80 % of illegally captured animals are birds, and 4 % of those are parrots. Although many seized birds do not survive, those that are recovered may be returned to the wild. The release of seized individuals into the wild should be conducted with caution, as local populations may suffer adverse effects if genetically different individuals are introduced. In this study, we evaluated the genetic relationships between 13 illegally captured blue-and-yellow macaws selected for release in northeastern Goiás, Brazil, and previously studied Brazilian macaw populations. We identified the seized macaws that were genetically similar to those from northwestern Goiás and that were therefore most suitable for release in that area. The genetic relationship was evaluated by sequence analysis of 403 bp of mitochondrial DNA control region. Relationships between mitochondrial haplotypes were computed via a median-joining network. Only six of the seized macaws were closely related to the macaws of northeastern Goiás, indicating that those macaws were potential candidates for release in that area. However, the release of these birds should follow all technical recommendations required by the Brazilian environmental authorities.     

Partner's behavior, not reward distribution, determines success in an unequal cooperative task in capuchin monkeys

It was recently demonstrated that capuchin monkeys notice and respond to distributional inequity, a trait that has been proposed to support the evolution of cooperation in the human species. However, it is unknown how capuchins react to inequitable rewards in an unrestricted cooperative paradigm in which they may freely choose both whether to participate and, within the bounds of their partner's behavior, which reward they will receive for their participation. We tested capuchin monkeys with such a design, using a cooperative barpull, which has been used with great success in the past. Contrary to our expectations, the equity of the reward distribution did not affect success or pulling behavior. However, the behavior of the partner in an unequal situation did affect overall success rates: pairs that had a tendency to alternate which individual received the higher-value food in unequal reward situations were more than twice as successful in obtaining rewards than pairs in which one individual dominated the higher-value food. This ability to equitably distribute rewards in inherently biased cooperative situations has profound implications for activities such as group hunts, in which multiple individuals work together for a single, monopolizable reward.     

An Investigation into the Cognition Behind Spontaneous String Pulling in New Caledonian Crows

The ability of some bird species to pull up meat hung on a string is a famous example of spontaneous animal problem solving. The “insight” hypothesis claims that this complex behaviour is based on cognitive abilities such as mental scenario building and imagination. An operant conditioning account, in contrast, would claim that this spontaneity is due to each action in string pulling being reinforced by the meat moving closer and remaining closer to the bird on the perch. We presented experienced and naïve New Caledonian crows with a novel, visually restricted string-pulling problem that reduced the quality of visual feedback during string pulling. Experienced crows solved this problem with reduced efficiency and increased errors compared to their performance in standard string pulling. Naïve crows either failed or solved the problem by trial and error learning. However, when visual feedback was available via a mirror mounted next to the apparatus, two naïve crows were able to perform at the same level as the experienced group. Our results raise the possibility that spontaneous string pulling in New Caledonian crows may not be based on insight but on operant conditioning mediated by a perceptual-motor feedback cycle.     

Phylogenetic relationships and historical biogeography of neotropical parrots (Psittaciformes: Psittacidae: Arini) inferred from mitochondrial and nuclear DNA sequences

Previous hypotheses of phylogenetic relationships among Neotropical parrots were based on limited taxon sampling and lacked support for most internal nodes. In this study we increased the number of taxa (29 species belonging to 25 of the 30 genera) and gene sequences (6388 base pairs of RAG-1, cyt b, NADH2, ATPase 6, ATPase 8, COIII, 12S rDNA, and 16S rDNA) to obtain a stronger molecular phylogenetic hypothesis for this group of birds. Analyses of the combined gene sequences using maximum likelihood and Bayesian methods resulted in a well-supported phylogeny and indicated that amazons and allies are a sister clade to macaws, conures, and relatives, and these two clades are in turn a sister group to parrotlets. Key morphological and behavioral characters used in previous classifications were mapped on the molecular tree and were phylogenetically uninformative. We estimated divergence times of taxa using the molecular tree and Bayesian and penalized likelihood methods that allow for rate variation in DNA substitutions among sites and taxa. Our estimates suggest that the Neotropical parrots shared a common ancestor with Australian parrots 59 Mya (million of years ago; 95% credibility interval (CrI) 66, 51 Mya), well before Australia separated from Antarctica and South America, implying that ancestral parrots were widespread in Gondwanaland. Thus, the divergence of Australian and Neotropical parrots could be attributed to vicariance. The three major clades of Neotropical parrots originated about 50 Mya (95% CrI 57, 41 Mya), coinciding with periods of higher sea level when both Antarctica and South America were fragmented with transcontinental seaways, and likely isolated the ancestors of modern Neotropical parrots in different regions in these continents. The correspondence between major paleoenvironmental changes in South America and the diversification of genera in the clade of amazons and allies between 46 and 16 Mya suggests they diversified exclusively in South America. Conversely, ancestors of parrotlets and of macaws, conures, and allies may have been isolated in Antarctica and/or the southern cone of South America, and only dispersed out of these southern regions when climate cooled and Antarctica became ice-encrusted about 35 Mya. The subsequent radiation of macaws and their allies in South America beginning about 28 Mya (95% CrI 22, 35 Mya) coincides with the uplift of the Andes and the subsequent formation of dry, open grassland habitats that would have facilitated ecological speciation via niche expansion from forested habitats.     

Cooperative Problem Solving by Orangutans (Pongo pygmaeus)

A captive pair of subadult male orangutans (Pongo pygmaeus) performed a cooperative task without training. Both partners had to pull a handle simultaneously in order for each to get food. They also learned the importance of the partner at the apparatus to make a successful response. The requirements of the cooperative task appear to have been understood by the orangutans, much like chimpanzees (Pan troglodytes) in the same situation. In contrast, capuchins (Cebus apella) succeeded in the cooperative task with a limited understanding of the requirement of the task and without taking into account the partner's role. These results gives further support to the hypothesis of a proximity of cognitive processes between chimpanzees and orangutans (in contrast to monkeys) though orangutans have not been seen to hunt cooperatively in the wild.