Mirror-mediated string-pulling task in Eurasian jays (Garrulus glandarius).

Mirror tasks can be used to investigate whether animals can instrumentally use a mirror to solve problems and can understand the correspondence between reflections and the real objects they represent. Two bird species, a corvid (New Caledonian crow) and a parrot (African grey parrot), have demonstrated the ability to use mirrors instrumentally in mirror-mediated spatial locating tasks. However, they have not been challenged with a mirror-guided reaching task, which involves a more complex understanding of the mirror's properties. In the present study, a task approximating the mirror-guided reaching task used in primate studies was adapted for, and given to, a corvid species (Eurasian jay) using a horizontal string-pulling paradigm. Four birds learned to pull the correct string to retrieve a food reward when they could see the food directly, whereas none used the reflected information to accomplish the same objective. Based on these results, it cannot be concluded whether these birds understand the correspondence between the location of the reward and its reflected information, or if the relative lack of visual-perceptual motor feedback given by the setup interfered with their performance. This novel task is posited to be conceptually more difficult compared to mirror-mediated spatial locating tasks, and should be used in avian species that have previously been successful at using the mirror instrumentally. This would establish whether these species can still succeed at it, and thus whether the task does indeed pose additional cognitive demands.

Stepping up to the thermogradient plate: a data framework for predicting seed germination under climate change.

BACKGROUND AND AIMS: Seed germination is strongly influenced by environmental temperatures. With global temperatures predicted to rise, the timing of germination for thousands of plant species could change, leading to potential decreases in fitness and ecosystem-wide impacts. The thermogradient plate (TGP) is a powerful but underutilized research tool that tests germination under a broad range of constant and alternating temperatures, giving researchers the ability to predict germination characteristics using current and future climates. Previously, limitations surrounding experimental design and data analysis methods have discouraged its use in seed biology research. METHODS: Here, we have developed a freely available R script that uses TGP data to analyse seed germination responses to temperature. We illustrate this analysis framework using three example species: Wollemia nobilis, Callitris baileyi and Alectryon subdentatus. The script generates >40 germination indices including germination rates and final germination across each cell of the TGP. These indices are then used to populate generalized additive models and predict germination under current and future monthly maximum and minimum temperatures anywhere on the globe. KEY RESULTS: In our study species, modelled data were highly correlated with observed data, allowing confident predictions of monthly germination patterns for current and future climates. Wollemia nobilis germinated across a broad range of temperatures and was relatively unaffected by predicted future temperatures. In contrast, C. baileyi and A. subdentatus showed strong seasonal temperature responses, and the timing for peak germination was predicted to shift seasonally under future temperatures. CONCLUSIONS: Our experimental workflow is a leap forward in the analysis of TGP experiments, increasing its many potential benefits, thereby improving research predictions and providing substantial information to inform management and conservation of plant species globally.

Convergent evolution of complex cognition: Insights from the field of avian cognition into the study of self-awareness.

Pioneering research on avian behaviour and cognitive neuroscience have highlighted that avian species, mainly corvids and parrots, have a cognitive tool kit comparable with apes and other large-brained mammals, despite conspicuous differences in their neuroarchitecture. This cognitive tool kit is driven by convergent evolution, and consists of complex processes such as casual reasoning, behavioural flexibility, imagination, and prospection. Here, we review experimental studies in corvids and parrots that tested complex cognitive processes within this tool kit. We then provide experimental examples for the potential involvement of metacognitive skills in the expression of the cognitive tool kit. We further expand the discussion of cognitive and metacognitive abilities in avian species, suggesting that an integrated assessment of these processes, together with revised and multiple tasks of mirror self-recognition, might shed light on one of the most highly debated topics in the literature-self-awareness in animals. Comparing the use of multiple assessments of self-awareness within species and across taxa will provide a more informative, richer picture of the level of consciousness in different organisms.

Transfer of physical understanding in a non-tool-using parrot.

Physical cognition has generally been assessed in tool-using species that possess a relatively large brain size, such as corvids and apes. Parrots, like corvids and apes, also have large relative brain sizes, yet although parrots rarely use tools in the wild, growing evidence suggests comparable performances on physical cognition tasks. It is, however, unclear whether success on such tasks is facilitated by previous experience and training procedures. We therefore investigated physical comprehension of object relationships in two non-tool-using species of captive neotropical parrots on a new means-end paradigm, the Trap-Gaps task, using unfamiliar materials and modified training procedures that precluded procedural cues. Red-shouldered macaws (Diopsittaca nobilis) and black-headed caiques (Pionites melanocephala) were presented with an initial task that required them to discriminate between pulling food trays through gaps while attending to the respective width of the gaps and size of the trays. Subjects were then presented with a novel, but functionally equivalent, transfer task. Six of eight birds solved the initial task through trial-and-error learning. Four of these six birds solved the transfer task, with one caique demonstrating spontaneous comprehension. These findings suggest that non-tool-using parrots may possess capacities for sophisticated physical cognition by generalising previously learned rules across novel problems.

An avian perspective on simulating other minds.

An exciting new study on ravens by Bugnyar, Reber, and Buckner (2016) raises important questions about whether nonhuman animals are capable of simulating other minds, rather than theorizing about them.

Role of facial familiarity and emotional expression intensity in ensemble emotion perception.

When looking at groups of people, we can extract information from the different faces to derive properties of the group, such as its average facial emotion, although how this average is computed remains a matter of debate. Here, we examined whether our participants' personal familiarity with the faces in the group, as well as the intensity of the facial expressions, biased ensemble perception. Participants judged the average emotional expression of ensembles of four different identities whose expressions depicted either neutral, angry, or happy emotions. For the angry and happy expressions, the intensity of the emotion could be either low (e.g., slightly happy) or high (very happy). When all the identities in the ensemble were unfamiliar, the presence of any high intensity emotional face biased ensemble perception towards its emotion. However, when a familiar face was present in the ensemble, perception was biased towards the familiar face's emotion regardless of its intensity. These findings reveal that how we perceive the average emotion of a group is influenced by both the emotional intensity and familiarity of the faces comprising the group, supporting the idea that different faces may be weighted differently in ensemble perception. These findings have important implications for the judgements we make about a group's overall emotional state may be biased by individuals within the group.

Soil chemistry and fungal communities are associated with dieback in an Endangered Australian shrub.

Background and aims: Field surveys across known populations of the Endangered Persoonia hirsuta (Proteaceae) in 2019 suggested the soil environment may be associated with dieback in this species. To explore how characteristics of the soil environment (e.g., pathogens, nutrients, soil microbes) relate to dieback, a soil bioassay (Experiment 1) was conducted using field soils from two dieback effected P. hirsuta populations. Additionally, a nitrogen addition experiment (Experiment 2) was conducted to explore how the addition of soil nitrogen impacts dieback. Methods: The field soils were baited for pathogens, and soil physiochemical and microbial community characteristics were assessed and related to dieback among plants in the field and nursery-grown plants inoculated with the same field soils. Roots from inoculated plants were harvested to confirm the presence of soil pathogens and root-associated endophytes. Using these isolates, a dual culture antagonism assay was performed to examine competition among these microbes and identify candidate pathogens or pathogen antagonists. Results: Dieback among plants in the field and Experiment 1 was associated with soil physiochemical properties (nitrogen and potassium), and soil microbes were identified as significant indicators of healthy and dieback-affected plants. Plants in Experiment 2 exhibited greater dieback when treated with elevated nitrogen. Additionally, post-harvest culturing identified fungi and other soil pathogens, some of which exhibited antagonistic behavior. Conclusion: This study identified candidate fungi and soil physiochemical properties associated with observed dieback and dieback resistance in an Endangered shrub and provides groundwork for further exploring what drives dieback and how it can be managed to promote the conservation of wild populations. Supplementary Information: The online version contains supplementary material available at 10.1007/s11104-022-05724-7.

Mirror stimulation in Eurasian jays (Garrulus glandarius).

Mirror exposure elicits a wide range of behavioral responses, some of which have been considered as part of possible evidence of mirror self-recognition (MSR). These responses can range from social behaviors, indicating that an animal considers its own reflection as a conspecific, to mirror-guided and self-directed actions. Evidence of MSR has been found categorically in only a few species, such as in magpies, chimpanzees, horses, and elephants. Evidence in corvids is currently debated due to inconsistent findings. In this study, we investigated the reaction of Eurasian jays when presenting them with three mirror-stimulation tasks. Based on the overall behavioral patterns across these three tasks, conclusions about birds' understanding of a reflective surface, and their perception of the reflection as either themselves or as a conspecific, appear premature. We highlight how the high neophobia of corvids and other methodological constraints might have hindered the likelihood to approach and explore a mirror, preventing the emergence of behaviors typically associated with MSR. Furthermore, we discuss how motivational factors, methodological constraints and species differences should be considered when interpreting behavioral responses to mirrors.

How does cognition evolve? Phylogenetic comparative psychology.

Now more than ever animal studies have the potential to test hypotheses regarding how cognition evolves. Comparative psychologists have developed new techniques to probe the cognitive mechanisms underlying animal behavior, and they have become increasingly skillful at adapting methodologies to test multiple species. Meanwhile, evolutionary biologists have generated quantitative approaches to investigate the phylogenetic distribution and function of phenotypic traits, including cognition. In particular, phylogenetic methods can quantitatively (1) test whether specific cognitive abilities are correlated with life history (e.g., lifespan), morphology (e.g., brain size), or socio-ecological variables (e.g., social system), (2) measure how strongly phylogenetic relatedness predicts the distribution of cognitive skills across species, and (3) estimate the ancestral state of a given cognitive trait using measures of cognitive performance from extant species. Phylogenetic methods can also be used to guide the selection of species comparisons that offer the strongest tests of a priori predictions of cognitive evolutionary hypotheses (i.e., phylogenetic targeting). Here, we explain how an integration of comparative psychology and evolutionary biology will answer a host of questions regarding the phylogenetic distribution and history of cognitive traits, as well as the evolutionary processes that drove their evolution.

Insightful problem solving and creative tool modification by captive nontool-using rooks.

The ability to use tools has been suggested to indicate advanced physical cognition in animals. Here we show that rooks, a member of the corvid family that do not appear to use tools in the wild are capable of insightful problem solving related to sophisticated tool use, including spontaneously modifying and using a variety of tools, shaping hooks out of wire, and using a series of tools in a sequence to gain a reward. It is remarkable that a species that does not use tools in the wild appears to possess an understanding of tools rivaling habitual tool users such as New Caledonian crows and chimpanzees. Our findings suggest that the ability to represent tools may be a domain-general cognitive capacity rather than an adaptive specialization and questions the relationship between physical intelligence and wild tool use.

Investigation of mirror-self recognition in ravens (Corvus corax).

Large-brained birds, such as corvids and parrots, tend to fail tests for self-recognition (mirror self-recognition [MSR]), but the limited positive evidence for MSR in these species has been questioned due to methodological limitations. In the present study, we aimed to investigate MSR in ravens by performing three mirror tests: a mirror exposure test, a mirror preference test, and a mark test. Across all three tests, the ravens' behavior was not consistent with MSR. Three out of six ravens infrequently interacted with the mirror and the nonmirror surfaces. Two birds explored the mirror and occasionally displayed contingent behaviors. Finally, the ravens made very few social displays toward the mirror, suggesting that at this stage they did not treat their reflection as a conspecific. These findings, along with the current evidence available, raise further questions on the validity of relying on one test to establish self-recognition and call for the development of methods beyond mirror tests to explore self-recognition in nonhuman animals. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Postconflict third-party affiliation in rooks, Corvus frugilegus.

Conflict features in the lives of many animal species and induces social stress mediated by glucocorticoid hormones [1]. Postconflict affiliation, between former opponents (reconciliation) or between former opponents and a bystander (third-party affiliation), has been suggested as a behavioral mechanism for reducing such stress [2], but has been studied almost exclusively in primates [3]. As with many primates, several bird species live in social groups and form affiliative relationships [4]. Do these distantly related animals also use affiliative behavior to offset the costs of conflict? We studied postconflict affiliation in a captive group of rooks. Unlike polygamous primates, monogamous rooks did not reconcile with former opponents. However, we found clear evidence of third-party affiliation after conflicts. Both initiators and targets of aggression engaged in third-party affiliation with a social partner and employed a specific behavior, bill twining, during the postconflict period. Both former aggressors and uninvolved third parties initiated affiliative contacts. Despite the long history of evolutionary divergence, the pattern of third-party affiliation in rooks is strikingly similar to that observed in tolerant primate species. Furthermore, the absence of reconciliation in rooks makes sense in light of the species differences in social systems.

Rooks perceive support relations similar to six-month-old babies.

Some corvids have demonstrated cognitive abilities that rival or exceed those of the great apes; for example, tool use in New Caledonian crows, and social cognition, episodic-like memory and future planning in Western scrub-jays. Rooks appear to be able to solve novel tasks through causal reasoning rather than simple trial-and-error learning. Animals with certain expectations about how objects interact would be able to narrow the field of candidate causes substantially, because some causes are simply 'impossible'. Here we present evidence that rooks hold such expectations and appear to possess perceptual understanding of support relations similar to that demonstrated by human babies, which is more comprehensive than that of chimpanzees.

Investigating physical cognition in rooks, Corvus frugilegus.

Although animals (particularly tool-users) are capable of solving physical tasks in the laboratory , the degree to which they understand them in terms of their underlying physical forces is a matter of contention. Here, using a new paradigm, the two-trap tube task, we report the performance of non-tool-using rooks. In contrast to the low success rates of previous studies using trap-tube problems , seven out of eight rooks solved the initial task, and did so rapidly. Instead of the usual, conceptually flawed control, we used a series of novel transfer tasks to test for understanding. All seven transferred their solution across a change in stimuli. However, six out of seven were unable to transfer to two further tasks, which did not share any one visual constant. One female was able to solve these further transfer tasks. Her result is suggestive evidence that rooks are capable of sophisticated physical cognition, if not through an understanding of unobservable forces , perhaps through rule abstraction. Our results highlight the need to investigate cognitive mechanisms other than causal understanding in studying animal physical cognition.

Chimpanzees solve the trap problem when the confound of tool-use is removed.

The trap-tube problem is difficult for chimpanzees to solve; in several studies only 1 to 2 subjects learn the solution. The authors tested eight chimpanzees on a non-tool-using version of the problem to investigate whether the inclusion of a tool in previous tests of the trap problem may have masked the ability of chimpanzees to solve it. All eight learned to avoid the trap, in 40 to 100 trials. One transferred to two tasks that had no visual cue in common. The authors examined the performance of 15 chimpanzees on a new task in a 2 x 2 design: seven had experience on the two-trap box, eight had not; half of each group was tested with a tool, half without one. An ANOVA revealed a significant effect of tool-inclusion and experience (p < .05). Our results show that including a tool in the trap problem profoundly affects the ability of chimpanzees to solve it. With regard to what the chimpanzees had learned, the results support the notion that rather than using the available stimuli as arbitrary cues, the subjects had encoded information about functional properties.

Western scrub-jays conceal auditory information when competitors can hear but cannot see.

Western scrub-jays (Aphelocoma californica) engage in a variety of cache-protection strategies to reduce the chances of cache theft by conspecifics. Many of these strategies revolve around reducing visual information to potential thieves. This study aimed to determine whether the jays also reduce auditory information during caching. Each jay was given the opportunity to cache food in two trays, one of which was filled with small pebbles that made considerable noise when cached in ('noisy' tray), whereas the other one contained soil that made little detectable noise when cached in ('quiet' tray). When the jays could be heard, but not seen, by a competitor, they cached proportionally less food items in the 'noisy' substrate than when they cached alone in the room, or when they could be seen and heard by competitors. These results suggest that western scrub-jays know when to conceal auditory information, namely when a competitor cannot see but can hear the caching event.

The development of caching and object permanence in Western scrub-jays (Aphelocoma californica): which emerges first?

Recent studies on the food-caching behavior of corvids have revealed complex physical and social skills, yet little is known about the ontogeny of food caching in relation to the development of cognitive capacities. Piagetian object permanence is the understanding that objects continue to exist even when they are no longer visible. Here, the authors focus on Piagetian Stages 3 and 4, because they are hallmarks in the cognitive development of both young children and animals. Our aim is to determine in a food-caching corvid, the Western scrub-jay, whether (1) Piagetian Stage 4 competence and tentative caching (i.e., hiding an item invisibly and retrieving it without delay), emerge concomitantly or consecutively; (2) whether experiencing the reappearance of hidden objects enhances the timing of the appearance of object permanence; and (3) discuss how the development of object permanence is related to behavioral development and sensorimotor intelligence. Our findings suggest that object permanence Stage 4 emerges before tentative caching, and independent of environmental influences, but that once the birds have developed simple object-permanence, then social learning might advance the interval after which tentative caching commences.

The Role of Emotional Expression and Eccentricity on Gaze Perception.

The perception of another's gaze direction and facial expression complements verbal communication and modulates how we interact with other people. However, our perception of these two cues is not always accurate, even when we are looking directly at the person. In addition, in many cases social communication occurs within groups of people where we can't always look directly at every person in the group. Here, we sought to examine how the presence of other people influences our perception of a target face. We asked participants to judge the direction of gaze of the target face as either looking to their left, to their right or directly at them, when the face was viewed on its own or viewed within a group of other identity faces. The target face either had an angry or a neutral expression and was viewed directly (foveal experiment), or within peripheral vision (peripheral experiment). When the target was viewed within a group, the flanking faces also had either neutral or angry expressions and their gaze was in one of five different directions (from averted leftwards to averted rightwards in steps of 10°). When the target face was viewed foveally there was no effect of target emotion on participants' judgments of its gaze direction. There was also no effect of the presence of flankers (regardless of expression) on the perception of the target gaze. When the target face was viewed peripherally, participants judged its direction of gaze to be direct over a wider range of gaze deviations than when viewed foveally, and more so for angry faces than neutral faces. We also find that flankers (regardless of emotional expression) did not influence performance. This suggests that observers judge that angry faces were looking at them over a broad range of gaze deviations in the periphery only, possibly resulting from increased uncertainty about the stimulus.

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.

Bilateral neurotoxic amygdala lesions in rhesus monkeys (Macaca mulatta): consistent pattern of behavior across different social contexts.

Although the amygdala has been repeatedly implicated in normal primate social behavior, great variability exists in the specific social and nonsocial behavioral changes observed in nonhuman primates with bilateral amygdala lesions. One plausible explanation pertains to differences in social context. This study measured the social behavior of amygdala-lesioned and unoperated rhesus monkeys (Macaca mulatta) in 2 contexts. Monkeys interacted in 4-member social groups over 32 test days. They were previously assessed in pairs (N. J. Emery et al., 2001) and were therefore familiar with each other at the beginning of this study. Across the 2 contexts, amygdala lesions produced a highly consistent pattern of social behavior. Operated monkeys engaged in more affiliative social interactions with control partners than did controls. In the course of their interactions, amygdala-lesioned monkeys also displayed an earlier decrease in nervous and fearful personality qualities than did controls. The increased exploration and sexual behavior recorded for amygdala-lesioned monkeys in pairs was not found in the 4-member groups. The authors concluded that the amygdala contributes to social inhibition and that this function transcends various social contexts.