Testing social learning of anti-predator responses in juvenile jackdaws: the importance of accounting for levels of agitation

This is the author accepted manuscript. The final version is available from the publisher via the DOI in this record.Social learning is often assumed to help young animals respond appropriately to potential threats in the environment. We brought wild, juvenile jackdaws briefly into captivity to test whether short exposures to conspecific vocalizations are sufficient to promote anti-predator learning. Individuals were presented with one of two models-a stuffed fox representing a genuine threat, or a toy elephant simulating a novel predator. Following an initial baseline presentation, juveniles were trained by pairing models with either adult mobbing calls, indicating danger, or contact calls suggesting no danger. In a final test phase with no playbacks, birds appeared to have habituated to the elephant, regardless of training, but responses to the fox remained high throughout, suggesting juveniles already recognized it as a predator before the experiment began. Training with mobbing calls did seem to generate elevated escape responses, but this was likely to be a carry-over effect of the playback in the previous trial. Overall, we found little evidence for social learning. Instead, individuals' responses were mainly driven by their level of agitation immediately preceding each presentation. These results highlight the importance of accounting for agitation in studies of anti-predator learning, and whenever animals are held in captivity for short periods.G.E.M. and A.T. were supported by a BBSRC David Phillips Fellowship to A.T. (BB/H021817/1 and BB/H021817/2). V.E.L. was supported by a Natural Environment Research Council Studentship

Testing relationship recognition in wild jackdaws (Corvus monedula)

This is the author accepted manuscriptAccording to the social intelligence hypothesis, understanding the challenges faced by social animals is key to understanding the evolution of cognition. In structured social groups, recognising the relationships of others is often important for predicting the outcomes of interactions. Third-party relationship recognition has been widely investigated in primates, but studies of other species are limited. Furthermore, few studies test for third-party relationship recognition in the wild, where cognitive abilities are deployed in response to natural socio-ecological pressures. Here, we used playback experiments to investigate whether wild jackdaws (Corvus monedula) track changes in their own relationships and the relationships of others. Females were presented with ‘infidelity simulations’: playbacks of their male partner copulating with a neighbouring female, and their male neighbour copulating with another female, against a congruent control. Our results showed substantial inter-individual variation in responses, but females did not respond more strongly to infidelity playbacks, indicating that jackdaws may not attend and/or respond to relationship information in this experimental context. Our results highlight the need for further study of relationship recognition and other cognitive traits that facilitate group-living in the wild, particularly in non-primates and in a wider range of social systems.BBSRC David Phillips Fellowshi

Evidence for individual discrimination and numerical assessment in collective antipredator behaviour in wild jackdaws (Corvus monedula)

This is the final version. Available on open access from the Royal Society via the DOI in this recordData accessibility: Data available from the Dryad Digital Repository: https://dx.doi.org/10.5061/dryad.006bn4kCollective responses to threats occur throughout the animal kingdom but little is known about the cognitive processes underpinning them. Antipredator mobbing is one such response. Approaching a predator may be highly risky, but the individual risk declines and the likelihood of repelling the predator increases in larger mobbing groups. The ability to appraise the number of conspecifics involved in a mobbing event could therefore facilitate strategic decisions about whether to join. Mobs are commonly initiated by recruitment calls, which may provide valuable information to guide decision-making. We tested whether the number of wild jackdaws responding to recruitment calls was influenced by the number of callers. As predicted, playbacks simulating three or five callers tended to recruit more individuals than playbacks of one caller. Recruitment also substantially increased if recruits themselves produced calls. These results suggest that jackdaws use individual vocal discrimination to assess the number of conspecifics involved in initiating mobbing events, and use this information to guide their responses. Our results show support for the use of numerical assessment in antipredator mobbing responses and highlight the need for a greater understanding of the cognitive processes involved in collective behaviour.Biotechnology and Biological Sciences Research Council (BBSRC)Human Frontier Science ProgramUniversity of Exete

Social learning about dangerous people by wild jackdaws

This is the final version. Available on open access from the Royal Society via the DOI in this recordData accessibility Data and R scripts associated with this work are available via Figshare (doi:10.6084/m9.figshare.7828781)For animals that live alongside humans, people can present both an opportunity and a threat. Previous studies have shown that several species can learn to discriminate between individual people and assess risk based on prior experience. To avoid potentially costly encounters, it may also pay individuals to learn about dangerous people based on information from others. Social learning about anthropogenic threats is likely to be beneficial in habitats dominated by human activity, but experimental evidence is limited. Here, we tested whether wild jackdaws (Corvus monedula) use social learning to recognize dangerous people. Using a within-subjects design, we presented breeding jackdaws with an unfamiliar person near their nest, combined with conspecific alarm calls. Subjects that heard alarm calls showed a heightened fear response in subsequent encounters with the person compared to a control group, reducing their latency to return to the nest. This study provides important evidence that animals use social learning to assess the level of risk posed by individual humans.Natural Environment Research Council (NERC)La Région Auvergne-Rhône-AlpesBiotechnology and Biological Sciences Research Council (BBSRC

Caller characteristics influence recruitment to collective antipredator events in jackdaws

This is the author accepted manuscript. Available from Nature Publishing Group via the DOI in this record.Across the animal kingdom, examples abound of individuals coming together to repel external threats. When such collective actions are initiated by recruitment signals, individuals may benefit from being selective in whom they join, so the identity of the initiator may determine the magnitude of the group response. However, the role of signaller discrimination in coordinating group-level responses has yet to be tested. Here we show that in wild jackdaws, a colonial corvid species, collective responses to anti-predator recruitment calls are mediated by caller characteristics. In playbacks next to nestboxes, the calls of nestbox residents attracted most recruits, followed in turn by other colony members, non-colony members and rooks (a sympatric corvid). Playbacks in fields outside nestbox colonies, where the immediate threat to broods was lower, showed similar results, with highest recruitment to nearby colony members’ calls. Responses were further influenced by caller sex: calls from non-colony member females were less likely to elicit responsive scolding by recruits than other calls, potentially reflecting social rank associated with sex and colony membership. These results show that vocal discrimination mediates jackdaws’ collective responses and highlight the need for further research into the cognitive basis of collective actions in animal groups.Biotechnology and Biological Sciences Research Council (BBSRC)Natural Environment Research Council (NERC

Costs and benefits of social relationships in the collective motion of bird flocks

This is the author accepted manuscript. The final version is available from Nature Research via the DOI in this record.Supplementary Figs. 1–12 and Supplementary Tables 1–3 are available in the Supplementary Information. Raw images captured by one of the four cameras and the reconstructed birds’ 3D movement trajectories are provided in Supplementary Videos 1–6. Plain text files, each including bird ID number, position, time, velocity, acceleration and wingbeat frequency at every time step, are provided in Supplementary Data 1–7. A plain text file that includes mean wingbeat frequency, flight speed and local density (approximated by the number of neighbours within a distance of 5 m from the focal bird) for paired and unpaired birds in six flocks, as well as for birds flying alone, is provided in Supplementary Data 8. All data required to reproduce the results in this study are included in Supplementary Data 1–8. Supplementary Data and Supplementary Videos are available at https://figshare.com/s/c55eb82bab800571d25d.Current understanding of collective behaviour in nature is based largely on models that assume that identical agents obey the same interaction rules, but in reality interactions may be influenced by social relationships among group members. Here, we show that social relationships transform local interactions and collective dynamics. We tracked individuals’ three-dimensional trajectories within flocks of jackdaws, a species that forms lifelong pair-bonds. Reflecting this social system, we find that flocks contain internal sub-structure, with discrete pairs of individuals tied together by spring-like effective forces. Within flocks, paired birds interacted with fewer neighbours than unpaired birds and flapped their wings more slowly, which may result in energy savings. However, flocks with more paired birds had shorter correlation lengths, which is likely to inhibit efficient information transfer through the flock. Similar changes to group properties emerge naturally from a generic self-propelled particle model. These results reveal a critical tension between individual- and group-level benefits during collective behaviour in species with differentiated social relationships, and have major evolutionary and cognitive implications.Human Frontiers in Science Programm

Local interactions and their group-level consequences in flocking jackdaws

This is the author accepted manuscript. The final version is available from the Royal Society via the DOI in this recordData accessibility: Data and code are available from the Dryad Digital Repository: https://doi.org/10.5061/dryad.kb8js06As one of nature's most striking examples of collective behaviour, bird flocks have attracted extensive research. However, we still lack an understanding of the attractive and repulsive forces that govern interactions between individuals within flocks and how these forces influence neighbours' relative positions and ultimately determine the shape of flocks. We address these issues by analysing the three-dimensional movements of wild jackdaws (Corvus monedula) in flocks containing 2–338 individuals. We quantify the social interaction forces in large, airborne flocks and find that these forces are highly anisotropic. The long-range attraction in the direction perpendicular to the movement direction is stronger than that along it, and the short-range repulsion is generated mainly by turning rather than changing speed. We explain this phenomenon by considering wingbeat frequency and the change in kinetic and gravitational potential energy during flight, and find that changing the direction of movement is less energetically costly than adjusting speed for birds. Furthermore, our data show that collision avoidance by turning can alter local neighbour distributions and ultimately change the group shape. Our results illustrate the macroscopic consequences of anisotropic interaction forces in bird flocks, and help to draw links between group structure, local interactions and the biophysics of animal locomotion.Human Frontiers in Science Programm

Wild jackdaws respond to their partner's distress, but not with consolation

This is the final version. Available on open access from the Royal Society via the DOI in this recordData accessibility: Data, R scripts and supplementary videos associated with this study are available in the figshare repository https://doi.org/10.6084/m9.figshare.13026815.v1Individuals are expected to manage their social relationships to maximize fitness returns. For example, reports of some mammals and birds offering unsolicited affiliation to distressed social partners (commonly termed 'consolation') are argued to illustrate convergent evolution of prosocial traits across divergent taxa. However, most studies cannot discriminate between consolation and alternative explanations such as self-soothing. Crucially, no study that controls for key confounds has examined consolation in the wild, where individuals face more complex and dangerous environments than in captivity. Controlling for common confounds, we find that male jackdaws (Corvus monedula) respond to their mate's stress-states, but not with consolation. Instead, they tended to decrease affiliation and partner visit rate in both experimental and natural contexts. This is striking because jackdaws have long-term monogamous relationships with highly interdependent fitness outcomes, which is precisely where theory predicts consolation should occur. Our findings challenge common conceptions about where consolation should evolve, and chime with concerns that current theory may be influenced by anthropomorphic expectations of how social relationships should be managed. To further our understanding of the evolution of such traits, we highlight the need for our current predictive frameworks to incorporate the behavioural trade-offs inherent to life in the wild.Natural Environment Research Council (NERC)Biotechnology and Biological Sciences Research Council (BBSRC)Leverhulme Trus

Trace elements in hemodialysis patients: a systematic review and meta-analysis

<p>Abstract</p> <p>Background</p> <p>Hemodialysis patients are at risk for deficiency of essential trace elements and excess of toxic trace elements, both of which can affect health. We conducted a systematic review to summarize existing literature on trace element status in hemodialysis patients.</p> <p>Methods</p> <p>All studies which reported relevant data for chronic hemodialysis patients and a healthy control population were eligible, regardless of language or publication status. We included studies which measured at least one of the following elements in whole blood, serum, or plasma: antimony, arsenic, boron, cadmium, chromium, cobalt, copper, fluorine, iodine, lead, manganese, mercury, molybdenum, nickel, selenium, tellurium, thallium, vanadium, and zinc. We calculated differences between hemodialysis patients and controls using the differences in mean trace element level, divided by the pooled standard deviation.</p> <p>Results</p> <p>We identified 128 eligible studies. Available data suggested that levels of cadmium, chromium, copper, lead, and vanadium were higher and that levels of selenium, zinc and manganese were lower in hemodialysis patients, compared with controls. Pooled standard mean differences exceeded 0.8 standard deviation units (a large difference) higher than controls for cadmium, chromium, vanadium, and lower than controls for selenium, zinc, and manganese. No studies reported data on antimony, iodine, tellurium, and thallium concentrations.</p> <p>Conclusion</p> <p>Average blood levels of biologically important trace elements were substantially different in hemodialysis patients, compared with healthy controls. Since both deficiency and excess of trace elements are potentially harmful yet amenable to therapy, the hypothesis that trace element status influences the risk of adverse clinical outcomes is worthy of investigation.</p

Nesting jackdaws’ responses to human voices vary with local disturbance levels and the gender of the speaker

This is the final version. Available from Elsevier via the DOI in this record. Data Availability: Data and R scripts used for this study are available in the figshare repository https://10.6084/m9.figshare.19249673.The ability to detect and respond to indicators of risk is vital for any animal and, for many species, humans represent a key threat. We investigated whether wild jackdaws, Corvus monedula, a species that thrives in anthropogenic environments but is regularly persecuted by people, associate human voices with differential degrees of risk and differ in their responses according to local levels of human disturbance. Playbacks showed that nesting females did not discriminate between the voices of familiar men who posed differing levels of threat, generalize to unfamiliar individuals with similar regional accents or discriminate between familiar and unfamiliar accents and voices. They were, however, considerably more wary towards male than female human voices, which may reflect the greater likelihood of negative experiences with men than women. Responses to playbacks also differed across fine-scale spatial locations: females nesting in areas of the colony with high levels of disturbance were less likely to leave the nest cup in response to playbacks and were more wary on their return to the nest than birds nesting in less disturbed areas. Nevertheless, levels of local disturbance did not influence reproductive success. Together these results indicate that, although vocal cues alone may not suffice for wild jackdaws to discriminate between individual humans or generalize across categories of people, sensitivity to cues of gender and local disturbance may help jackdaws to optimize their defensive behaviour and maintain breeding success. Further research into plastic responses towards indicators of human risk is vital to understand and mitigate the impacts of increasing urbanization on wildlife populations.Biotechnology and Biological Sciences Research Council (BBSRC)Natural Environment Research Council (NERC