In social animal groups, an individual's spatial position is a major determinant of both predation risk and foraging rewards. Additionally, the occupation of positions in the front of moving groups is generally assumed to correlate with the initiation of group movements. However, whether some individuals are predisposed to consistently occupy certain positions and, in some instances, to consistently lead groups over time is as yet unresolved in many species. Using the mosquitofish (Gambusia holbrooki), we examined the consistency of individuals' spatial positions within a moving group over successive trials. We found that certain individuals consistently occupied front positions in moving groups and also that it was typically these individuals that initiated group decisions. The number of individuals involved in leading the group varied according to the amount of information held by group members, with a greater number of changes in leadership in a novel compared to a relatively familiar environment. Finally, our results show that the occupation of lead positions in moving groups was not explained by characteristics such as dominance, size or sex, suggesting that certain individuals are predisposed to leadership roles. This suggests that being a leader or a follower may to some extent be an intrinsic property of the individual.
Despite the frequency with which mixed-species groups are observed in nature, studies of collective behaviour typically focus on single-species groups. Here, we quantify and compare the patterns of interactions between three fish species, threespine sticklebacks (Gasterosteus aculeatus), ninespine sticklebacks (Pungitius pungitius) and roach (Rutilus rutilus) in both single- and mixed-species shoals in the laboratory. Pilot data confirmed that the three species form both single- and mixed-species shoals in the wild. In our laboratory study, we found that single-species groups were more polarized than mixed-species groups, while single-species groups of threespine sticklebacks and roach were more cohesive than mixed shoals of these species. Furthermore, while there was no difference between the inter-individual distances between threespine and ninespine sticklebacks within mixed-species groups, there was some evidence of segregation by species in mixed groups of threespine sticklebacks and roach. There were differences between treatments in mean pairwise transfer entropy, and in particular we identify species-differences in information use within the mixed-species groups, and, similarly, differences in responses to conspecifics and heterospecifics in mixed-species groups. We speculate that differences in the patterns of interactions between species in mixed-species groups may determine patterns of fission and fusion in such groups.
Studies on the effects of visitors on zoo animals have shown mixed findings and as a result, the manner in which visitors affect zoo animals remains unclear for many species, including a rarely studied taxa such as penguins. Penguins are a common zoo-housed species and have been shown to display huddling, vigilance and avoidance towards zoo visitors which can be indicative of fear. Here, we examined the effects of covering one visitor viewing area window, out of four, on little penguin (Eudyptula minor) behaviours that may be indicative of fear. Two treatments were randomly imposed on different days: 1) The main visitor viewing area window, where most visitor-penguin interactions occurred, was uncovered (‘Main window uncovered’) and 2) The main visitor viewing area window was covered (‘Main window covered’). Penguin numbers and behaviour were recorded near the main visitor viewing area window and the three other visitor viewing area windows, as well as one area not visible to visitors (‘Corner’ area). Furthermore, visitor numbers and visitor behaviour were recorded at all four visitor viewing area windows. Covering the main visitor viewing area window reduced the proportion of visitors present at this window by about 85% (p < 0.001) and reduced potentially threatening visitor behaviours at this window such as tactile contact with the window, loud vocalisations and sudden movement (p < 0.05). When the main visitor viewing area window was covered, the proportion of penguins present increased by about 25% (p < 0.05), the proportion of visible penguins preening in the water increased by about 180% (p < 0.05) and the proportion of visible penguins vigilant decreased by about 70% (p < 0.05) in the area near this main window. A preference for the Corner area was also found whereby 59% and 49% of penguins were present in this area when the main window was uncovered and covered, respectively. These results provide limited evidence that the little penguins in this exhibit showed an aversion to the area near the main visitor viewing area window when it was uncovered based on the increased avoidance and vigilance and decreased preening in the water in this area. This suggests visitors may be fear-provoking for these little penguins. However, it is unclear whether visual contact with visitors per se or other aspects of visitor contact, such as visitor-induced sounds and vibrations, were responsible for this apparent aversion when this window was uncovered.
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