Abstract: Rich circumstantial evidence suggests that the extensive behavioural diversity recorded in wild great apes reflects a complexity of cultural variation unmatched by species other than our own. However, the capacity for cultural transmission assumed by this interpretation has remained difficult to test rigorously in the field, where the scope for controlled experimentation is limited. Here we show that experimentally introduced technologies will spread within different ape communities. Unobserved by group mates, we first trained a high-ranking female from each of two groups of captive chimpanzees to adopt one of two different tool-use techniques for obtaining food from the same 'Pan-pipe' apparatus, then re-introduced each female to her respective group. All but two of 32 chimpanzees mastered the new technique under the influence of their local expert, whereas none did so in a third population lacking an expert. Most chimpanzees adopted the method seeded in their group, and these traditions continued to diverge over time. A subset of chimpanzees that discovered the alternative method nevertheless went on to match the predominant approach of their companions, showing a conformity bias that is regarded as a hallmark of human culture.

Abstract: In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences.

Abstract: The role of dominant individuals in leading groups of animals is not yet well understood. We investigated whether dominant beef cows, Bos taurus, have more influence on herd movement on pasture than more subordinate cows. A herd of 15 Gasconne cows was observed for a 3-week period between dawn and dusk. The positions of all adult cows were recorded with GPS collars at 1 min intervals and the behaviour of each cow was recorded in 5 min scans. The dominance hierarchy was recorded by ad libitum sampling. Through cluster analysis of the recorded data, we distinguished three herd behaviour patterns: resting, foraging and travelling. Dominant cows were closer to the front of the herd during both travelling and foraging. During travelling, more dominant cows also had more direct trajectories and were more aligned both with their nearest neighbours and with the whole herd. During foraging, the trajectories of dominant cows were shorter than those of subordinate cows. The results indicate that foraging and short-distance travelling movements by female beef cattle are not led by any particular individual but rather are influenced by a graded type of leadership; that is, the more dominant a cow is, the stronger the influence it may have on the movements of the herd.

Abstract: Cardiac morphology in human athletes is known to differ, depending on the sports-specific endurance component of their events, whereas anecdotes abound about superlative athletes with large hearts. As the heart determines stroke volume and maximum O(2) uptake in mammals, we undertook a study to test the hypothesis that the morphology of the equine heart would differ between trained horses, depending on race type, and that left ventricular size would be greatest in elite performers. Echocardiography was performed in 482 race-fit Thoroughbreds engaged in either flat (1,000-2,500 m) or jump racing (3,200-6,400 m). Body weight and sex-adjusted measures of left ventricular size were largest in horses engaged in jump racing over fixed fences, compared with horses running shorter distances on the flat (range 8-16%). The observed differences in cardiac morphologies suggest that subtle differences in training and competition result in cardiac adaptations that are appropriate to the endurance component of the horses' event. Derived left ventricular mass was strongly associated with published rating (quality) in horses racing over longer distances in jump races (P < or = 0.001), but less so for horses in flat races. Rather, left ventricular ejection fraction and left ventricular mass combined were positively associated with race rating in older flat racehorses running over sprint (<1,408 m) and longer distances (>1,408 m), explaining 25-35% of overall variation in performance, as well as being closely associated with performance in longer races over jumps (23%). These data provide the first direct evidence that cardiac size influences athletic performance in a group of mammalian running athletes.

Abstract: The classic evolutionary theory of aging explains why mortality rises with age: as individuals grow older, less lifetime fertility remains, so continued survival contributes less to reproductive fitness. However, successful reproduction often involves intergenerational transfers as well as fertility. In the formal theory offered here, age-specific selective pressure on mortality depends on a weighted average of remaining fertility (the classic effect) and remaining intergenerational transfers to be made to others. For species at the optimal quantity-investment tradeoff for offspring, only the transfer effect shapes mortality, explaining postreproductive survival and why juvenile mortality declines with age. It also explains the evolution of lower fertility, longer life, and increased investments in offspring.