Temporal and Contextual Consistency of Leadership in Homing Pigeon Flocks

Organized flight of homing pigeons (Columba livia) was previously shown to rely on simple leadership rules between flock mates, yet the stability of this social structuring over time and across different contexts remains unclear. We quantified the repeatability of leadership-based flock structures within a flight and across multiple flights conducted with the same animals. We compared two contexts of flock composition: flocks of birds of the same age and flight experience; and, flocks of birds of different ages and flight experience. All flocks displayed consistent leadership-based structures over time, showing that individuals have stable roles in the navigational decisions of the flock. However, flocks of balanced age and flight experience exhibited reduced leadership stability, indicating that these factors promote flock structuring. Our study empirically demonstrates that leadership and followership are consistent behaviours in homing pigeon flocks, but such consistency is affected by the heterogeneity of individual flight experiences and/or age. Similar evidence from other species suggests leadership as an important mechanism for coordinated motion in small groups of animals with strong social bonds.     

Aerial Surveys for Estimating Wild Turkey Abundance in the Texas Rolling Plains

Abstract: Aerial surveys have been used to estimate abundance of several wild bird species including wild turkeys (Meleagris gallopavo). We used inflatable turkey decoys at 3 study sites in the Texas Rolling Plains to simulate Rio Grande wild turkey (M. g. intermedia) flocks. We evaluated detectability of flocks and errors in counting flock size during fixed-wing (Cessna 172) aerial surveys using logistic and linear regression models. Flock detectability was primarily influenced by flock size and vegetative cover, and errors in counting flock size were primarily influenced by size of flocks. We conducted computer simulations to evaluate the accuracy and precision of fixed-wing aerial surveys and examined power to detect trends in population change. Our simulations suggested abundance estimates from fixed-wing aerial surveys may be underestimated by 10-15% (2.0-4.8% CV). Power analyses suggested that fixed-wing aerial surveys can provide sufficient power (>0.80) to detect a population change of 10-25% over a 4-5-year period. We concluded fixed-wing aerial surveys are feasible on ecoregion scales.     

Does mixed-species flocking influence how birds respond to a gradient of land-use intensity?

Conservation biology is increasingly concerned with preserving interactions among species such as mutualisms in landscapes facing anthropogenic change. We investigated how one kind of mutualism, mixed-species bird flocks, influences the way in which birds respond to different habitat types of varying land-use intensity. We use data from a well-replicated, large-scale study in Sri Lanka and the Western Ghats of India, in which flocks were observed inside forest reserves, in ‘buffer zones'' of degraded forest or timber plantations, and in areas of intensive agriculture. We find flocks affected the responses of birds in three ways: (i) species with high propensity to flock were more sensitive to land use; (ii) different flock types, dominated by different flock leaders, varied in their sensitivity to land use and because following species have distinct preferences for leaders, this can have a cascading effect on followers'' habitat selection; and (iii) those forest-interior species that remain outside of forests were found more inside flocks than would be expected by chance, as they may use flocks more in suboptimal habitat. We conclude that designing policies to protect flocks and their leading species may be an effective way to conserve multiple bird species in mixed forest and agricultural landscapes.     

Decision accuracy in complex environments is often maximized by small group sizes

Individuals in groups, whether composed of humans or other animal species, often make important decisions collectively, including avoiding predators, selecting a direction in which to migrate and electing political leaders. Theoretical and empirical work suggests that collective decisions can be more accurate than individual decisions, a phenomenon known as the ‘wisdom of crowds’. In these previous studies, it has been assumed that individuals make independent estimates based on a single environmental cue. In the real world, however, most cues exhibit some spatial and temporal correlation, and consequently, the sensory information that near neighbours detect will also be, to some degree, correlated. Furthermore, it may be rare for an environment to contain only a single informative cue, with multiple cues being the norm. We demonstrate, using two simple models, that taking this natural complexity into account considerably alters the relationship between group size and decision-making accuracy. In only a minority of environments do we observe the typical wisdom of crowds phenomenon (whereby collective accuracy increases monotonically with group size). When the wisdom of crowds is not observed, we find that a finite, and often small, group size maximizes decision accuracy. We reveal that, counterintuitively, it is the noise inherent in these small groups that enhances their accuracy, allowing individuals in such groups to avoid the detrimental effects of correlated information while exploiting the benefits of collective decision-making. Our results demonstrate that the conventional view of the wisdom of crowds may not be informative in complex and realistic environments, and that being in small groups can maximize decision accuracy across many contexts.     

Epidemiological Characteristics of Classical Scrapie Outbreaks in 30 Sheep Flocks in the United Kingdom

Background

Most previous analyses of scrapie outbreaks have focused on flocks run by research institutes, which may not reflect the field situation. Within this study, we attempt to rectify this deficit by describing the epidemiological characteristics of 30 sheep flocks naturally-infected with classical scrapie, and by exploring possible underlying causes of variation in the characteristics between flocks, including flock-level prion protein (PrP) genotype profile. In total, the study involved PrP genotype data for nearly 8600 animals and over 400 scrapie cases.

Methodology/Principal Findings

We found that most scrapie cases were restricted to just two PrP genotypes (ARQ/VRQ and VRQ/VRQ), though two flocks had markedly different affected genotypes, despite having similar underlying genotype profiles to other flocks of the same breed; we identified differences amongst flocks in the age of cases of certain PrP genotypes; we found that the age-at-onset of clinical signs depended on peak incidence and flock type; we found evidence that purchasing infected animals is an important means of introducing scrapie to a flock; we found some evidence that flock-level PrP genotype profile and flock size account for variation in outbreak characteristics; identified seasonality in cases associated with lambing time in certain flocks; and we identified one case that was homozygous for phenylalanine at codon 141, a polymorphism associated with a very high risk of atypical scrapie, and 28 cases that were heterozygous at this codon.

Conclusions/Significance

This paper presents the largest study to date on commercially-run sheep flocks naturally-infected with classical scrapie, involving 30 study flocks, more than 400 scrapie cases and over 8500 PrP genotypes. We show that some of the observed variation in epidemiological characteristics between farms is related to differences in their PrP genotype profile; although much remains unexplained and may instead be attributed to the stochastic nature of scrapie dynamics.     

Habitat use of Tibetan Eared Pheasant Crossoptilon harmani flocks in the non-breeding season

Habitat use by Tibetan Eared Pheasant Crossoptilon harmani flocks in shrub vegetation was investigated in the Lhasa area of Tibet during the non-breeding season of 19951996. Home range composition varied considerably among flocks, but stream belts were consistently used as foraging grounds. Slope direction, altitude and vegetation had little effect on habitat selection. In the absence of supplemental food, core range size was positively correlated with flock size, suggesting that food supplementation could support larger flocks. Flocks regularly roosted on the ground at midday at two or three relatively fixed sites within core ranges. At night they used patches of relatively tall, dense vegetation at the year-round sites in areas near cliffs or in hollows. The size of the night-roost site was related to flock size. Our results strongly suggested that both foraging and night-roosting habitats in the shrub environment are crucial to the birds.     

Vigilance in Greater Flamingos Wintering in Southern Tunisia: Age-Dependent Flock Size Effect

Decrease in individual vigilance with flock size is a widely recognized pattern in group‐living species. However such a relationship may be affected by other factors, such as age and flock composition. For instance, because young animals generally lack experience and have higher nutritional needs than adults, they can be expected not only to be less vigilant than adults but also to decrease their vigilance level by a greater extent when flock size increases than adults do. We investigated this issue using data on greater flamingos wintering in the gulf of Gabès, in southern Tunisia. Flamingos tended to congregate in small single‐age flocks for feeding, but as flock size increased, flocks became mixed. We found that when flock size increased, young flamingos significantly decreased their vigilance time, while adult did not, suggesting an age‐dependent flock size effect on vigilance. However, when flock composition (single‐age vs. mixed) was taken into account, a more complex pattern was found. Within single‐age and small flocks, no difference was found between young flamingos and adult ones regarding their vigilance level and their response to increasing flock size. However, within mixed and large flocks, adult flamingos were more vigilant than young ones, while variation in flock size did not result in a significant change in vigilance. These results suggest that young birds relied on the presence of adults, and hence more experienced individuals in detecting dangers, to reduce their vigilance and to increase their foraging time in order to satisfy their higher nutritional requirements. They could also be interpreted as a possible consequence of increasing competition with flock size which constrained more nutritionally stressed young flamingos to increase their foraging time to the detriment of vigilance.     

How do vigilance and feeding by common cranes Grus grus depend on age,habitat, and flock size?

Animals often spend less time vigilant and more time feeding when foraging in larger groups. This group-size effect does not, however, consider if larger groups differ systematically from smaller ones: Large groups could form in different habitats than small groups or be composed of a different mix of ages or classes than small groups. We examined how habitat differences and flock size and composition explain feeding and vigilance rates in common cranes Grus grus , wintering in holm oak Quercus ilex dehesas of Spain. Flock size and composition were related to habitat type in cranes: flocks formed in areas sown with cereal crops were larger than flocks formed in set aside areas. Vigilance rate depended on habitat but decreased with increasing flock size in a similar way across all habitats. Juveniles were less vigilant than adults and showed little change in vigilance with flock size. Vigilance increased and feeding time decreased over months from November through February. Our results show that vigilance is affected by habitat but that the group size effect on vigilance is not the product of differences between habitats in group size or composition.     

Number of neighbors instead of group size significantly affects individual vigilance levels in large animal aggregations

The group size effect states that animals living in groups gain anti‐predator benefits through reducing vigilance levels as group size increases. A basic assumption of group size effect is that all individuals are equally important for a focal individual, who may adjust its vigilance levels according to social information acquired from them. However, some studies have indicated that neighbors pose greater influences on an individual's vigilance decisions than other group members, especially in large aggregations. Vigilance has also been found to be directed to both predators (anti‐predation vigilance) and conspecifics (social vigilance). Central individuals might rely more on social vigilance than peripheral individuals. To test these hypotheses, we examined the effects of flock size, number of neighbors and position within a flock on vigilance and competition of greater white‐fronted goose Anser albifrons that form large foraging flocks in winter, controlling the effects of other variables (group identity, winter period and site). We found that individual vigilance levels were significantly affected by number of neighbors and position within a flock, whereas flock size showed no effect. Individuals devoted a large component of vigilance to nearby flock mates. Central individuals directed a relatively larger proportion of vigilance to monitor neighbors than peripheral ones, indicating that central individuals more relied on social information acquired from neighbors, possibly caused by the more blocked visual field of central individuals. Moreover, some social vigilance may function as conducting or preventing agonistic interactions since competition intensity was positively correlated with number of neighbors. Our study therefore demonstrate that the number of neighbors is more important than group size in determining individual vigilance in large animal groups. Further studies are still needed to unravel which neighbors pose greater influence on individual vigilance, and the factors that influence individuals to acquire information from their neighbors to adjust vigilance behaviors.     

Relation of group size and daily activity patterns to southern lapwing (Vanellus chilensis) behaviour

Behavioural patterns of birds commonly vary according to flock size and daily activity pattern. Southern lapwing behaviours and their relation with flock size were studied, as well as the relationship between the frequency of behaviours and the period of the day. Results showed that the proportion of time spent in foraging and vigilance was higher during the morning, when small groups were more common, and decreased from midday on, when group size increased. Maintenance and inactivity (sleeping) behaviours presented the opposite pattern. Correlation between flock size and period of the day, and their similar effects on bird behaviour may be evidence that groups of different sizes could have different functions throughout the day. Our results show that southern lapwings seem to form smaller flocks for feeding in the beginning of the day and larger flocks later for different activities (e.g., maintenance and sleeping). In this sense, it is possible that group size variation throughout the day is related to different demands for specific behaviours (functions) of groups, according to the daily activity patterns of the birds.     

暂时与群体分离的个体藏马鸡的反捕食警戒

通过与同伴分担捕食风险 ,生活在稳定群体中的个体动物可以获得长期的适合度利益 ,但同时它们不得不承担食物竞争所带来的潜在代价。这种代价常常取决于食物资源的类型。当好的食物资源出现时 ,一些个体可以离开群体而独享这种资源。了解这些临时游离者如何组织其反捕食行为 ,在进化生态学上是有意义的。藏马鸡 (Crossoptilonharmani)是西藏雅鲁藏布江中游高山灌丛植被的一种典型的非繁殖季节集群鸟类。野外观察表明 ,为了独享好的食物资源 ,一些个体常常远离当前群体的活动范围。分离事件更可能发生于大的群体 ,但其发生率与参与者的数量呈负相关 ;而参与分离的个体愈多 ,分离持续的时间就愈长。分离者的个体警惕水平随着临时群体大小的增加而下降 ,遵从在其它自然大小鸟类群体所发现的一般性规律。分离行为的发生和持续时间被认为是个体对当前食物回报和捕食风险进行权衡的结果。这种利益 -代价权衡也可以解释藏马鸡所具有的强烈集群行为     

Continuous decisions

Humans and other animals evolved to make decisions that extend over time with continuous and ever-changing options. Nonetheless, the academic study of decision-making is mostly limited to the simple case of choice between two options. Here, we advocate that the study of choice should expand to include continuous decisions. Continuous decisions, by our definition, involve a continuum of possible responses and take place over an extended period of time during which the response is continuously subject to modification. In most continuous decisions, the range of options can fluctuate and is affected by recent responses, making consideration of reciprocal feedback between choices and the environment essential. The study of continuous decisions raises new questions, such as how abstract processes of valuation and comparison are co-implemented with action planning and execution, how we simulate the large number of possible futures our choices lead to, and how our brains employ hierarchical structure to make choices more efficiently. While microeconomic theory has proven invaluable for discrete decisions, we propose that engineering control theory may serve as a better foundation for continuous ones. And while the concept of value has proven foundational for discrete decisions, goal states and policies may prove more useful for continuous ones.This article is part of the theme issue ‘Existence and prevalence of economic behaviours among non-human primates’.     

Are vocal characteristics related to leadership patterns in mixed‐species bird flocks?

What structures the organization of mixed‐species bird flocks, so that some ‘nuclear’ species lead the flocks, and others follow? Previous research has shown that species actively listen to each other, and that leaders are gregarious; such gregarious species tend to make contact calls and hence may be vocally conspicuous. Here we investigated whether vocal characteristics are associated with leadership, using a global dataset of mixed‐species flock studies and recordings from sound archives. We first asked whether leaders are different from following or occasional species in flocks in the proportion of the recordings that contain calls (n = 58 flock studies, 145 species), and especially alarm calls (n = 111 species). We found that leaders tended to have a higher proportion of their vocalizations that were classified as calls than occasional species, and both leaders and following species had a significantly higher proportion of their calls rated as alarms compared to occasional species. Next, we investigated the acoustic characteristics of flock participants’ calls, hypothesizing that leaders would make more calls, and have less silence on the recordings. We also hypothesized that leaders’ calls would be simple acoustically, as contact calls tend to be, and thus similar to each other, as well as being detectable, in being low frequency and with high frequence bandwidth. The analysis (n = 45 species, 169 recordings) found that only one of these predictions was supported: leading species were less often silent than following or occasional species. Unexpectedly, leaders’ calls were less similar to each other than occasional species. The greater amount of information available and the greater variety of that information support the hypothesis that leadership in flocks is related to vocal communication. We highlight the use of sound archives to ask questions about behavioral and community ecology, while acknowledging some limitations of such studies.     

Group decisions and individual differences: route fidelity predicts flight leadership in homing pigeons (Columba livia)

How social-living animals make collective decisions is currently the subject of intense scientific interest, with increasing focus on the role of individual variation within the group. Previously, we demonstrated that during paired flight in homing pigeons, a fully transitive leadership hierarchy emerges as birds are forced to choose between their own and their partner''s habitual routes. This stable hierarchy suggests a role for individual differences mediating leadership decisions within homing pigeon pairs. What these differences are, however, has remained elusive. Using novel quantitative techniques to analyse habitual route structure, we show here that leadership can be predicted from prior route-following fidelity. Birds that are more faithful to their own route when homing alone are more likely to emerge as leaders when homing socially. We discuss how this fidelity may relate to the leadership phenomenon, and propose that leadership may emerge from the interplay between individual route confidence and the dynamics of paired flight.     

Individual-level personality influences social foraging and collective behaviour in wild birds

There is increasing evidence that animal groups can maintain coordinated behaviour and make collective decisions based on simple interaction rules. Effective collective action may be further facilitated by individual variation within groups, particularly through leader–follower polymorphisms. Recent studies have suggested that individual-level personality traits influence the degree to which individuals use social information, are attracted to conspecifics, or act as leaders/followers. However, evidence is equivocal and largely limited to laboratory studies. We use an automated data-collection system to conduct an experiment testing the relationship between personality and collective decision-making in the wild. First, we report that foraging flocks of great tits (Parus major) show strikingly synchronous behaviour. A predictive model of collective decision-making replicates patterns well, suggesting simple interaction rules are sufficient to explain the observed social behaviour. Second, within groups, individuals with more reactive personalities behave more collectively, moving to within-flock areas of higher density. By contrast, proactive individuals tend to move to and feed at spatial periphery of flocks. Finally, comparing alternative simulations of flocking with empirical data, we demonstrate that variation in personality promotes within-patch movement while maintaining group cohesion. Our results illustrate the importance of incorporating individual variability in models of social behaviour.     

Self-organization of collective escape in pigeon flocks

Bird flocks under predation demonstrate complex patterns of collective escape. These patterns may emerge by self-organization from local interactions among group-members. Computational models have been shown to be valuable for identifying what behavioral rules may govern such interactions among individuals during collective motion. However, our knowledge of such rules for collective escape is limited by the lack of quantitative data on bird flocks under predation in the field. In the present study, we analyze the first GPS trajectories of pigeons in airborne flocks attacked by a robotic falcon in order to build a species-specific model of collective escape. We use our model to examine a recently identified distance-dependent pattern of collective behavior: the closer the prey is to the predator, the higher the frequency with which flock members turn away from it. We first extract from the empirical data of pigeon flocks the characteristics of their shape and internal structure (bearing angle and distance to nearest neighbors). Combining these with information on their coordination from the literature, we build an agent-based model adjusted to pigeons’ collective escape. We show that the pattern of turning away from the predator with increased frequency when the predator is closer arises without prey prioritizing escape when the predator is near. Instead, it emerges through self-organization from a behavioral rule to avoid the predator independently of their distance to it. During this self-organization process, we show how flock members increase their consensus over which direction to escape and turn collectively as the predator gets closer. Our results suggest that coordination among flock members, combined with simple escape rules, reduces the cognitive costs of tracking the predator while flocking. Such escape rules that are independent of the distance to the predator can now be investigated in other species. Our study showcases the important role of computational models in the interpretation of empirical findings of collective behavior.     

The ecological significance of extremely large flocks of birds

Population size is generally limited by resource availability during and outside the breeding season. Therefore, maximum size of flocks may provide important information on population regulation and the influence of diet and trophic level on maximal degree of sociality. We hypothesized that (a) flock size should increase with nutrient availability; (b) flock size should decrease with latitude because productivity is higher at lower latitude; (c) aquatic habitats should have larger flocks than terrestrial habitats because the former are less accessible; (d) smaller species should have larger flocks because they require overall less food; (e) human‐impacted species that live in perturbed habitats should have smaller flocks than other species; (f) flock size should decrease with increasing trophic level because there is a reduction in biomass due to conversion at each trophic level; and (g) flocks of species depending on ancestral landscapes should have decreased in size in recent years due to human impact (e.g., land‐use). We obtained 1564 observations of flocks that exceeded 100,000 individuals in order to test the predictions listed above. Most effect sizes were small to medium accounting for 1%–9% of the variance, while large effects accounting for 25% or more were only found for total nitrogen used per km² and area used for agriculture. Changes in large bird flocks were caused by habitat degradation and persecution, and temporal decline in size of large flocks revealed changes in nutrient use, reductions in nutrient cycling, and changes in flock size linked to trophic level.     

Investigation of a Simple Model for Within-Flock Transmission of Scrapie

Genetic control programs for scrapie in sheep build on solid knowledge of how susceptibility to scrapie is modulated by the prion protein genotype at the level of an individual sheep. In order to satisfactorily analyze the effectivity of control programs at the population level, insight is needed at the flock level, i.e., how the grouping of sheep in flocks affects the population-level transmission risk. In particular, one would like to understand how this risk is affected by between-flock differences in genotype frequency distribution. A first step is to model the scrapie transmission risk within a flock as a function of the flock genotype profile. Here we do so by estimating parameters for a model of within-flock transmission using genotyping data on Dutch flocks affected by scrapie. We show that the data are consistent with a relatively simple transmission model assuming horizontal transmission and homogeneous mixing between animals. The model expresses the basic reproduction number for within-flock scrapie as a weighted average of genotype-specific susceptibilities, multiplied by a single overall transmission parameter. The value of the overall transmission parameter may vary between flocks to account for random between-flock variation in non-genetic determinants such as management practice. Here we provide an estimate of its mean value and variation for Dutch flocks.