Intra- and interspecific variation in self-control capacities of parrots in a delay of gratification task.

Forgoing immediate satisfaction for higher pay-offs in the future (delayed gratification) could be adaptive in situations that wild animals may encounter. To explain species-differences in self-control, hypotheses based on social complexity, feeding ecology, brain size and metabolic rate have been proposed. To explore these hypotheses in a comparative setting, we tested three macaw species (neotropical parrots)-great green macaws (N = 8), blue-throated macaws (N = 6), blue-headed macaws (N = 6)-and the distantly related African grey parrots (afrotropical parrots; N = 8) in a modified rotating tray task, in which subjects are required to inhibit consuming a constantly available low-quality reward in favour of a high-quality reward that becomes available only after an increasing delay (min. 5 s, max. 60 s). All four species successfully waited for a minimum of 8.3 s ± 11.7 s (group level mean ± SD) with African greys reaching a delay of 29.4 ± 15.2 s, and great green macaws-as best performing macaw species-tolerating delays of 20 s ± 8 s. The best performing African grey individual reached a maximum delay of 50 s, whereas, a great green and a blue-throated macaw tolerated a delay of 30 s max. Females tolerated higher maximum delays than males. Engaging in distraction behaviours enhanced waiting performance across species and all birds were able to anticipate the waiting duration. Our results suggest that both feeding and socio-ecological complexity may be a factor in self-control, but further systematically collected comparative data on self-control of different (parrot) species are required to test the evolutionary hypotheses rigorously.

Social interaction analysis in captive orcas (Orcinus orca).

The management of socially complex species in captivity is challenging. Research on their social behavior improves our understanding of interactions in captive animals and captive-group management. We conducted a detailed analysis of social relationships shown by the orcas kept at Loro Parque zoo and their tendency to reconcile after aggressive episodes. Affiliative interactions were the most frequent social activities compared to agonistic or sexual interactions. Within affiliative behaviors, we documented the pattern "gentle tongue bite", where an animal touches the other's tongue with his teeth but does not bite it. Affiliative interactions between a specific pair of orcas occurred significantly more often than expected by chance, and together with low levels of agonistic interactions, indicated particular affinity between some individuals. The most frequently observed low-level agonistic relationship was that of the two older males (Tekoa-Keto); however, they also showed frequent sexual and affiliative interactions. Sexual-like behaviors (pursuit, mount, and penis between males) were found in both sexes. Finally, the observed corrected conciliatory tendency (31.57%) was within the range described for other primate and cetacean species. This study provides a systematic way to assess social interactions as well as conflict management strategies in cetaceans housed in zoos and zoo-like facilities and may help to improve animal welfare and management of animals in controlled environments.

Are parrots poor at motor self-regulation or is the cylinder task poor at measuring it?

The ability to inhibit unproductive motor responses triggered by salient stimuli is a fundamental inhibitory skill. Such motor self-regulation is thought to underlie more complex cognitive mechanisms, like self-control. Recently, a large-scale study, comparing 36 species, found that absolute brain size best predicted competence in motor inhibition, with great apes as the best performers. This was challenged when three Corvus species (corvids) were found to parallel great apes despite having much smaller absolute brain sizes. However, new analyses suggest that it is the number of pallial neurons, and not absolute brain size per se, that correlates with levels of motor inhibition. Both studies used the cylinder task, a detour-reaching test where food is presented behind a transparent barrier. We tested four species from the order Psittaciformes (parrots) on this task. Like corvids, many parrots have relatively large brains, high numbers of pallial neurons, and solve challenging cognitive tasks. Nonetheless, parrots performed markedly worse than the Corvus species in the cylinder task and exhibited strong learning effects in performance and response times. Our results suggest either that parrots are poor at controlling their motor impulses, and hence that pallial neuronal numbers do not always correlate with such skills, or that the widely used cylinder task may not be a good measure of motor inhibition.

Testing problem-solving capacities: differences between individual testing and social group setting.

Testing animals individually in problem-solving tasks limits distractions of the subjects during the test, so that they can fully concentrate on the problem. However, such individual performance may not indicate the problem-solving capacity that is commonly employed in the wild when individuals are faced with a novel problem in their social groups, where the presence of a conspecific influences an individual's behaviour. To assess the validity of data gathered from parrots when tested individually, we compared the performance on patterned-string tasks among parrots tested singly and parrots tested in social context. We tested two captive groups of orange-winged amazons (Amazona amazonica) with several patterned-string tasks. Despite the differences in the testing environment (singly vs. social context), parrots from both groups performed similarly. However, we found that the willingness to participate in the tasks was significantly higher for the individuals tested in social context. The study provides further evidence for the crucial influence of social context on individual's response to a challenging situation such as a problem-solving test.

Means-end comprehension in four parrot species: explained by social complexity.

A comparative approach is required to investigate the evolutionary origins of cognitive abilities. In this paper, we compare the performance of four parrot species, spectacled parrotlets (Forpus conspicillatus), rainbow lorikeets (Trichoglossus haematodus), green-winged macaws (Ara chloroptera) and sulphur-crested cockatoos (Cacatua galerita triton) in standardized string-pulling and string-choice paradigms. We varied the spatial relationship between the strings, the presence of a reward and the physical contact between the string and the reward to test different cognitive skills requiring means-end comprehension. The species tested showed a high individual and inter-specific variation in their ability to solve the tasks. Spectacled parrotlets performed best among the four species and solved the most complex choice tasks, namely crossed-string task and broken-string task, spontaneously. In contrast, macaws and cockatoos failed to identify the correct string in these two tasks. The rainbow lorikeets were outperformed by the parrotlets, but outperformed in turn the macaws and the cockatoos. The findings can be best explained by the variation in social complexity among species, rather than in their ecology.

Comparative cognition: Practical shortcomings and some potential ways forward.

The objectives in the field of comparative cognition are clear; efforts are devoted to revealing the selection pressures that shape the brains and cognitive abilities of different species and understanding cognitive processes in differently structured brains. However, our progress on reaching these objectives is slow, mostly because of several major practical challenges. In this review, we discuss 2 major shortcomings: (a) the poor systematics and low magnitude of the phylogenetic comparisons made, and (b) the weak comparability of the results caused by interfering species-specific confounding factors (perceptual, motivational, and morphological) alongside an insufficient level of standardisation of the methodologies. We propose a multiple-level comparative approach that emphasises the importance of achieving more direct comparisons within taxonomic groups at genus or family level as the first step before comparing between distantly related groups. We also encourage increasing interdisciplinary efforts to execute "team-science" approach in building a systematic and direct large-scale phylogenetic comparisons of bigger cognitive test batteries that produce reliable species-representative data. We finally revisit some existing suggestions that allow us to maximise standardisation while minimising species-specific confounding factors. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Assessing African grey parrots' prosocial tendencies in a token choice paradigm.

Prosociality is defined as a voluntary, typically low-cost behaviour that benefits another individual. Social tolerance has been proposed as a potential driver for its evolution, both on the proximate and on the ultimate level. Parrots are an interesting species to study such other-regarding behaviours, given that they are highly social and stand out in terms of relative brain size and cognitive capacity. We tested eight African grey parrots in a dyadic prosocial choice test. They faced a choice between two different tokens, a prosocial (actor and partner rewarded) and a selfish (only actor rewarded) one. We found that the birds did not behave prosocially when one subject remained in the actor role; however, when roles were alternated, the birds' prosocial choices increased. The birds also seemed to reciprocate their partner's choices, given that a contingency between choices was observed. If the food provisioned to the partner was of higher quality than that the actor obtained, actors increased their willingness to provide food to their partner. Nonetheless, the control conditions suggest that the parrots did not fully understand the task's contingencies. In sum, African grey parrots show the potential for prosociality and reciprocity; however, considering their lack of understanding of the contingencies of the particular tasks used in this study, the underlying motivation for the observed behaviour remains to be addressed by future studies, in order to elucidate the phylogenetic distribution of prosociality further.

Parrots do not show inequity aversion.

Inequity aversion, the negative reaction to unequal treatment, is considered a mechanism for stabilizing cooperative interactions between non-kin group members. However, this might only be adaptive for species that switch cooperative partners. Utilizing a comparative approach, inequity aversion has been assessed in many mammalian species and recently also in corvids and one parrot species, kea, revealing mixed results. To broaden our knowledge about the phylogenetic distribution of inequity aversion, we tested four parrot species in the token exchange paradigm. We varied the quality of rewards delivered to dyads of birds, as well as the effort required to obtain a reward. Blue-headed macaws and African grey parrots showed no reaction to being rewarded unequally. The bigger macaws were less willing to exchange tokens in the "unequal" condition compared to the "equal high" condition in which both birds obtained high quality rewards, but a closer examination of the results and the findings from the control conditions reveal that inequity aversion does not account for it. None of the species responded to inequity in terms of effort. Parrots may not exhibit inequity aversion due to interdependence on their life-long partner and the high costs associated with finding a new partner.

Economic Decision-Making in Parrots.

Economic decision-making involves weighing up differently beneficial alternatives to maximise payoff. This sometimes requires the ability to forego one's desire for immediate satisfaction. This ability is considered cognitively challenging because it not only requires inhibiting impulses, but also evaluating expected outcomes in order to decide whether waiting is worthwhile. We tested four parrot species in a token exchange task. The subjects were first trained to exchange three types of tokens for a food item of low, medium, and high value and successfully learned to exchange these in an order according to their value. Subsequently, they were confronted with a choice between a food item and a token that could be exchanged for higher-quality food. In additional control conditions however, choosing a token led to an equal or lower payoff. Individuals of all species were capable of deciding economically, yet only large macaws outperformed the other species in one of the crucial controls. For some individuals, particularly African grey parrots, the token apparently had an intrinsic value, which prevented them from choosing economically in some control conditions and which should be considered as potentially confounding by future token exchange studies.

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.