Significance Cooperation in animals is an enigma because it contravenes the basic notion that evolution favors selfish genes that promote only their own well-being. Bird migration in organized ...V-shaped or echelon formations constitutes such a cooperation dilemma. We show that juvenile Northern bald ibis ( Geronticus eremita ) cooperate by taking turns and precisely matching times they spend in the advantageous trailing position and in the disadvantageous front position. This time matching is done on a pairwise level. Furthermore, we found evidence that the animals' propensity to reciprocate in leading has a substantial influence on the size and cohesion of the flight formations. This study shows that direct reciprocation can enable cooperation between animals in a natural context.
One conspicuous feature of several larger bird species is their annual migration in V-shaped or echelon formation. When birds are flying in these formations, energy savings can be achieved by using the aerodynamic up-wash produced by the preceding bird. As the leading bird in a formation cannot profit from this up-wash, a social dilemma arises around the question of who is going to fly in front? To investigate how this dilemma is solved, we studied the flight behavior of a flock of juvenile Northern bald ibis ( Geronticus eremita ) during a human-guided autumn migration. We could show that the amount of time a bird is leading a formation is strongly correlated with the time it can itself profit from flying in the wake of another bird. On the dyadic level, birds match the time they spend in the wake of each other by frequent pairwise switches of the leading position. Taken together, these results suggest that bald ibis cooperate by directly taking turns in leading a formation. On the proximate level, we propose that it is mainly the high number of iterations and the immediacy of reciprocation opportunities that favor direct reciprocation. Finally, we found evidence that the animals' propensity to reciprocate in leading has a substantial influence on the size and cohesion of the flight formations.
Continuing degradation of coral reef ecosystems has generated substantial interest in how management can support reef resilience. Fishing is the primary source of diminished reef function globally, ...leading to widespread calls for additional marine reserves to recover fish biomass and restore key ecosystem functions. Yet there are no established baselines for determining when these conservation objectives have been met or whether alternative management strategies provide similar ecosystem benefits. Here we establish empirical conservation benchmarks and fish biomass recovery timelines against which coral reefs can be assessed and managed by studying the recovery potential of more than 800 coral reefs along an exploitation gradient. We show that resident reef fish biomass in the absence of fishing (B0) averages ∼1,000 kg ha(-1), and that the vast majority (83%) of fished reefs are missing more than half their expected biomass, with severe consequences for key ecosystem functions such as predation. Given protection from fishing, reef fish biomass has the potential to recover within 35 years on average and less than 60 years when heavily depleted. Notably, alternative fisheries restrictions are largely (64%) successful at maintaining biomass above 50% of B0, sustaining key functions such as herbivory. Our results demonstrate that crucial ecosystem functions can be maintained through a range of fisheries restrictions, allowing coral reef managers to develop recovery plans that meet conservation and livelihood objectives in areas where marine reserves are not socially or politically feasible solutions.
Cooperative hunting is believed to have important implications for the evolution of sociality and advanced cognitive abilities. Variation in the level of hunt organisation amongst species and how ...their cognitive, behavioural and athletic adaptations may contribute to observed patterns of cooperative hunting behaviour, however, are poorly understood. We, therefore, reviewed the literature for evidence of different levels of hunt organisation and cooperation in carnivorans and examined their social and physical adaptations for hunting. Descriptions of group hunting were scarce for many species and often of insufficient detail for us to be able to classify the level of hunt organisation involved. However, despite this, reports of behaviour fitting the description of collaboration, the most advanced level of hunt organisation, were found in over half the carnivorans reported to hunt cooperatively. There was no evidence that this behaviour would require advanced cognitive abilities. However, there was some evidence that both social mechanisms reducing aggression between group members and information transfer amongst individuals may aid cooperative hunting. In general, the cooperative strategies used seemed to depend partly on the species' locomotor abilities and habitat. There was some evidence that individuals take on consistent roles during cooperative hunts in some species, but it was not clear if this reflects individuals' physical differences, social factors or life experiences. Better understanding of the social, cognitive and physical mechanisms underlying cooperative hunting, and indeed establishing to what degree it exists in the first instance, will require more data for multiple individuals and species over many hunts.
Giraffes (
) possess specialised anatomy. Their disproportionately elongate limbs and neck confer recognised feeding advantages, but little is known about how their morphology affects locomotor ...function. In this study, we examined the stride parameters and ground reaction forces from three adult giraffes in a zoological park, across a range of walking speeds. The patterns of GRFs during walking indicate that giraffes, similar to other mammalian quadrupeds, maintain a forelimb-biased weight distribution. The angular excursion of the neck has functional links with locomotor dynamics in giraffes, and was exaggerated at faster speeds. The horizontal accelerations of the neck and trunk were out of phase compared with the vertical accelerations, which were intermediate between in and out of phase. Despite possessing specialised morphology, giraffes' stride parameters were broadly predicted from dynamic similarity, facilitating the use of other quadrupedal locomotion models to generate testable hypotheses in giraffes.
Herding of sheep by dogs is a powerful example of one individual causing many unwilling individuals to move in the same direction. Similar phenomena are central to crowd control, cleaning the ...environment and other engineering problems. Despite single dogs solving this ‘shepherding problem’ every day, it remains unknown which algorithm they employ or whether a general algorithm exists for shepherding. Here, we demonstrate such an algorithm, based on adaptive switching between collecting the agents when they are too dispersed and driving them once they are aggregated. Our algorithm reproduces key features of empirical data collected from sheep–dog interactions and suggests new ways in which robots can be designed to influence movements of living and artificial agents.
The cheetah and racing greyhound are of a similar size and gross morphology and yet the cheetah is able to achieve a far higher top speed. We compared the kinematics and kinetics of galloping in the ...cheetah and greyhound to investigate how the cheetah can attain such remarkable maximum speeds. This also presented an opportunity to investigate some of the potential limits to maximum running speed in quadrupeds, which remain poorly understood. By combining force plate and high speed video data of galloping cheetahs and greyhounds, we show how the cheetah uses a lower stride frequency/longer stride length than the greyhound at any given speed. In some trials, the cheetahs used swing times as low as those of the greyhounds (0.2 s) so the cheetah has scope to use higher stride frequencies (up to 4.0 Hz), which may contribute to it having a higher top speed that the greyhound. Weight distribution between the animal's limbs varied with increasing speed. At high speed, the hindlimbs support the majority of the animal's body weight, with the cheetah supporting 70% of its body weight on its hindlimbs at 18 m s(-1); however, the greyhound hindlimbs support just 62% of its body weight. Supporting a greater proportion of body weight on a particular limb is likely to reduce the risk of slipping during propulsive efforts. Our results demonstrate several features of galloping and highlight differences between the cheetah and greyhound that may account for the cheetah's faster maximum speeds.
1. Evaluating landscape connectivity and identifying and protecting corridors for animal movement have become central challenges in applied ecology and conservation. Currently, resource selection ...analyses are widely used to focus corridor planning where animal movement is predicted to occur. An animal's behavioural state (e.g. foraging, dispersing) is a significant determinant of resource selection patterns, yet has largely been ignored in connectivity assessments. 2. We review 16 years of connectivity studies employing resource selection analysis to evaluate how researchers have incorporated animal behaviour into corridor planning, and highlight promising new approaches for identifying wildlife corridors. To illustrate the importance of behavioural information in such analyses, we present an empirical case study to test behaviour-specific predictions of connectivity with long-distance dispersal movements of African wild dogs Lycaon pictus. We conclude by recommending strategies for developing more realistic connectivity models for future conservation efforts. 3. Our review indicates that most connectivity studies conflate resource selection with connectivity requirements, which may result in misleading estimates of landscape resistance, and lack validation of proposed connectivity models with movement data. 4. Our case study shows that including only directed movement behaviour when measuring resource selection reveals markedly different, and more accurate, connectivity estimates than a model measuring resource selection independent of behavioural state. 5. Synthesis and applications. Our results, using African wild dogs as a case study, suggest that resource selection analyses that fail to consider an animal's behavioural state may be insufficient in targeting movement pathways and corridors for protection. This failure may result in misidentification of wildlife corridors and misallocation of limited conservation resources. Our findings underscore the need for considering patterns of animal movement in appropriate behavioural contexts to ensure the effective application of resource selection analyses for corridor planning.
The fastest and most manoeuvrable terrestrial animals are found in savannah habitats, where predators chase and capture running prey. Hunt outcome and success rate are critical to survival, so both ...predator and prey should evolve to be faster and/or more manoeuvrable. Here we compare locomotor characteristics in two pursuit predator-prey pairs, lion-zebra and cheetah-impala, in their natural savannah habitat in Botswana. We show that although cheetahs and impalas were universally more athletic than lions and zebras in terms of speed, acceleration and turning, within each predator-prey pair, the predators had 20% higher muscle fibre power than prey, 37% greater acceleration and 72% greater deceleration capacity than their prey. We simulated hunt dynamics with these data and showed that hunts at lower speeds enable prey to use their maximum manoeuvring capacity and favour prey survival, and that the predator needs to be more athletic than its prey to sustain a viable success rate.
Many species travel in highly organized groups. The most quoted function of these configurations is to reduce energy expenditure and enhance locomotor performance of individuals in the assemblage. ...The distinctive V formation of bird flocks has long intrigued researchers and continues to attract both scientific and popular attention. The well-held belief is that such aggregations give an energetic benefit for those birds that are flying behind and to one side of another bird through using the regions of upwash generated by the wings of the preceding bird, although a definitive account of the aerodynamic implications of these formations has remained elusive. Here we show that individuals of northern bald ibises (Geronticus eremita) flying in a V flock position themselves in aerodynamically optimum positions, in that they agree with theoretical aerodynamic predictions. Furthermore, we demonstrate that birds show wingtip path coherence when flying in V positions, flapping spatially in phase and thus enabling upwash capture to be maximized throughout the entire flap cycle. In contrast, when birds fly immediately behind another bird--in a streamwise position--there is no wingtip path coherence; the wing-beats are in spatial anti-phase. This could potentially reduce the adverse effects of downwash for the following bird. These aerodynamic accomplishments were previously not thought possible for birds because of the complex flight dynamics and sensory feedback that would be required to perform such a feat. We conclude that the intricate mechanisms involved in V formation flight indicate awareness of the spatial wake structures of nearby flock-mates, and remarkable ability either to sense or predict it. We suggest that birds in V formation have phasing strategies to cope with the dynamic wakes produced by flapping wings.
Coral reefs provide ecosystem goods and services for millions of people in the tropics, but reef conditions are declining worldwide. Effective solutions to the crisis facing coral reefs depend in ...part on understanding the context under which different types of conservation benefits can be maximized. Our global analysis of nearly 1,800 tropical reefs reveals how the intensity of human impacts in the surrounding seascape, measured as a function of human population size and accessibility to reefs (“gravity”), diminishes the effectiveness of marine reserves at sustaining reef fish biomass and the presence of top predators, even where compliance with reserve rules is high. Critically, fish biomass in high-compliance marine reserves located where human impacts were intensive tended to be less than a quarter that of reserves where human impacts were low. Similarly, the probability of encountering top predators on reefs with high human impacts was close to zero, even in high-compliance marine reserves. However, we find that the relative difference between openly fished sites and reserves (what we refer to as conservation gains) are highest for fish biomass (excluding predators) where human impacts are moderate and for top predators where human impacts are low. Our results illustrate critical ecological trade-offs in meeting key conservation objectives: reserves placed where there are moderate-to-high human impacts can provide substantial conservation gains for fish biomass, yet they are unlikely to support key ecosystem functions like higher-order predation, which is more prevalent in reserve locations with low human impacts.
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