"Leading according to need" in self-organizing groups.

Self-organizing-system approaches have shed significant light on the mechanisms underlying synchronized movements by large groups of animals, such as shoals of fish, flocks of birds, or herds of ungulates. However, these approaches rarely consider conflicts of interest between group members, although there is reason to suppose that such conflicts are commonplace. Here, we demonstrate that, where conflicts exist, individual members of self-organizing groups can, in principle, increase their influence on group movement destination by strategically changing simple behavioral parameters (namely, movement speed, assertiveness, and social attraction range). However, they do so at the expense of an increased risk of group fragmentation and a decrease in movement efficiency. We argue that the resulting trade-offs faced by each group member render it likely that group movements are led by those members for which reaching a particular destination is most crucial or group cohesion is least important. We term this phenomenon leading according to "need" or "social indifference," respectively. Both kinds of leading can occur in the absence of knowledge of or communication about the needs of other group members and without the assumption of altruistic cooperation. We discuss our findings in the light of observations on fish and other vertebrates.

Survey of badger access to farm buildings and facilities in relation to contact with cattle.

Thirty-six farms in parishes in western England that had recently experienced herd breakdowns of bovine tuberculosis were surveyed for signs of badger activity and for husbandry practices relating to the access of badgers to the farm buildings and facilities. Signs of activity were detected within the farmyards and buildings of 14 of the farms and were associated with water troughs at pasture on two of them. Few of the farmers implemented practices to reduce contact between badgers and cattle. Stored cattle feed was freely accessible to wild animals in 88 per cent of the feed stores. Two badger carcases, and two of 66 samples of badger droppings, cultured positive for Mycobacterium bovis. Signs of badgers within farmyards were significantly positively associated with the number of badger setts and latrines in the immediate vicinity, but were not related to any recorded farm husbandry procedures.

Democracy in animals: the evolution of shared group decisions.

A 'consensus decision' is when the members of a group choose, collectively, between mutually exclusive actions. In humans, consensus decisions are often made democratically or in an 'equally shared' manner, i.e. all group members contribute to the decision. Biologists are only now realizing that shared consensus decisions also occur in social animals (other than eusocial insects). Sharing of decisions is, in principle, more profitable for groups than accepting the 'unshared' decision of a single dominant member. However, this is not true for all individual group members, posing a question as to how shared decision making could evolve. Here, we use a game theory model to show that sharing of decisions can evolve under a wide range of circumstances but especially in the following ones: when groups are heterogeneous in composition; when alternative decision outcomes differ in potential costs and these costs are large; when grouping benefits are marginal; or when groups are close to, or above, optimal size. Since these conditions are common in nature, it is easy to see how mechanisms for shared decision making could have arisen in a wide range of species, including early human ancestors.

Nonrandom movement behavior at habitat boundaries in two butterfly species: implications for dispersal.

We observed meadow brown (Maniola jurtina) and gatekeeper (Pyronia tithonus) butterflies at habitat boundaries and observed spontaneous movements out of suitable habitat in order to investigate such movements in relation to dispersal. We found that butterflies of both species were aware of the position of a highly permeable habitat boundary without needing to cross it. Nevertheless, a considerable proportion of butterflies close to the boundary left their habitat (25-43%). Butterflies that crossed the boundary, and moved substantial distances into unsuitable habitat (up to 350 m in M. jurtina and 70 m in P. tithonus), usually returned to their original habitat patch (98-100%). Movement trajectories, at least in M. jurtina, were significantly different from, and more directed and systematic than, a correlated random walk. Approximately 70-80% of spontaneous movements into unsuitable habitat in both species were "foray" loops comparable to those described in mammals and birds. We conclude that, since migrants seemed to have considerable control over leaving their patch and over their subsequent movement trajectories, chance encounter rates with habitat boundaries, and indeed habitat leaving rates, might be less crucial in determining dispersal rates than is usually assumed. In addition, random dispersal trajectories should not be taken for granted in population or evolution models.

Energetics reveals physiologically distinct castes in a eusocial mammal.

Eusociality, which occurs among mammals only in two species of African mole-rat, is characterized by division of labour between morphologically distinct 'castes'. In Damaraland mole-rats (Cryptomys damarensis), colony labour is divided between 'infrequent worker' and 'frequent worker' castes. Frequent workers are active year-round and together perform more than 95% of the total work of the colony, whereas infrequent workers typically perform less than 5% of the total work. Anecdotal evidence suggests that infrequent workers may act as dispersers, with dispersal being limited to comparatively rare periods when the soil is softened by moisture. Here we show that infrequent workers and queens increase their daily energy expenditure after rainfall whereas frequent workers do not. Infrequent workers are also fatter than frequent workers. We suggest that infrequent workers constitute a physiologically distinct dispersing caste, the members of which, instead of contributing to the work of the colony and helping the queen to reproduce, build up their own body reserves in preparation for dispersal and reproduction when environmental conditions are suitable.

Foray search: an effective systematic dispersal strategy in fragmented landscapes.

In the absence of evidence to the contrary, population models generally assume that the dispersal trajectories of animals are random, but systematic dispersal could be more efficient at detecting new habitat and may therefore constitute a more realistic assumption. Here, we investigate, by means of simulations, the properties of a potentially widespread systematic dispersal strategy termed "foray search." Foray search was more efficient in detecting suitable habitat than was random dispersal in most landscapes and was less subject to energetic constraints. However, it also resulted in considerably shorter net dispersed distances and higher mortality per net dispersed distance than did random dispersal, and it would therefore be likely to lead to lower dispersal rates toward the margins of population networks. Consequently, the use of foray search by dispersers could crucially affect the extinction-colonization balance of metapopulations and the evolution of dispersal rates. We conclude that population models need to take the dispersal trajectories of individuals into account in order to make reliable predictions.

Reliable microsatellite genotyping of the Eurasian badger (Meles meles) using faecal DNA.

The potential link between badgers and bovine tuberculosis has made it vital to develop accurate techniques to census badgers. Here we investigate the potential of using genetic profiles obtained from faecal DNA as a basis for population size estimation. After trialling several methods we obtained a high amplification success rate (89%) by storing faeces in 70% ethanol and using the guanidine thiocyanate/silica method for extraction. Using 70% ethanol as a storage agent had the advantage of it being an antiseptic. In order to obtain reliable genotypes with fewer amplification reactions than the standard multiple-tubes approach, we devised a comparative approach in which genetic profiles were compared and replication directed at similar, but not identical, genotypes. This modified method achieved a reduction in polymerase chain reactions comparable with the maximum-likelihood model when just using reliability criteria, and was slightly better when using reliability criteria with the additional proviso that alleles must be observed twice to be considered reliable. Our comparative approach would be best suited for studies that include multiple faeces from each individual. We utilized our approach in a well-studied population of badgers from which individuals had been sampled and reliable genotypes obtained. In a study of 53 faeces sampled from three social groups over 10 days, we found that direct enumeration could not be used to estimate population size, but that the application of mark-recapture models has the potential to provide more accurate results.

Group decision-making in animals.

Groups of animals often need to make communal decisions, for example about which activities to perform, when to perform them and which direction to travel in; however, little is known about how they do so. Here, we model the fitness consequences of two possible decision-making mechanisms: 'despotism' and 'democracy'. We show that under most conditions, the costs to subordinate group members, and to the group as a whole, are considerably higher for despotic than for democratic decisions. Even when the despot is the most experienced group member, it only pays other members to accept its decision when group size is small and the difference in information is large. Democratic decisions are more beneficial primarily because they tend to produce less extreme decisions, rather than because each individual has an influence on the decision per se. Our model suggests that democracy should be widespread and makes quantitative, testable predictions about group decision-making in non-humans.

Use of cattle farm resources by badgers (Meles meles) and risk of bovine tuberculosis (Mycobacterium bovis) transmission to cattle.

Nocturnal observations, radio telemetry and time-lapse camera surveillance were used to investigate visits by badgers (Meles meles L.) to two cattle farms. During 59 half-nights (ca. 295 h) of observation and 17 nights (ca. 154 h) of camera surveillance, 139 separate visits to farm buildings, by at least 26 individually identifiable badgers from two social groups, were recorded. The badgers, which included three individuals infected with bovine tuberculosis (Mycobacterium bovis), used cowsheds, feedsheds, barns, haystacks, slurry pits, cattle troughs and farmyards to exploit a range of food resources, including cattle feed and silage. Cattle feed was contaminated with badger faeces and badgers also came into close contact with cattle. The minimum number of badgers visiting farm buildings per night was negatively correlated with local 24 h rainfall. We conclude that exploitation by badgers of resources provided by cattle farms constitutes a potentially important mechanism for tuberculosis transmission from badgers to cattle.

Non-random dispersal in the butterfly Maniola jurtina: implications for metapopulation models.

The dispersal patterns of animals are important in metapopulation ecology because they affect the dynamics and survival of populations. Theoretical models assume random dispersal but little is known in practice about the dispersal behaviour of individual animals or the strategy by which dispersers locate distant habitat patches. In the present study, we released individual meadow brown butterflies (Maniola jurtina) in a non-habitat and investigated their ability to return to a suitable habitat. The results provided three reasons for supposing that meadow brown butterflies do not seek habitat by means of random flight. First, when released within the range of their normal dispersal distances, the butterflies orientated towards suitable habitat at a higher rate than expected at random. Second, when released at larger distances from their habitat, they used a non-random, systematic, search strategy in which they flew in loops around the release point and returned periodically to it. Third, butterflies returned to a familiar habitat patch rather than a non-familiar one when given a choice. If dispersers actively orientate towards or search systematically for distant habitat, this may be problematic for existing metapopulation models, including models of the evolution of dispersal rates in metapopulations.

Activity synchrony and social cohesion: a fission-fusion model.

A social group can only be spatially coherent if its members synchronize activities such as foraging and resting. However, activity synchronization is costly to individuals if it requires them to postpone an activity that would be personally more profitable in order to do what the rest of the group is doing. Such costs will be particularly high in groups whose members belong to different age, size or sex classes since the optimal allocation of time to various activities is likely to differ between such classes. Thus, differences in the costs of activity synchronization between and within classes could cause non-homogenous groups to be less stable than homogenous groups, with the result that homogenous groups predominate in the population: that is, they could cause 'social segregation' of animals of different sex, size or age. We develop a model that predicts the degree of social segregation attributable to differences in activity synchronization between homogenous and non-homogenous groups and use this model in determining whether activity synchronization can explain intersexual social segregation in red deer (Cervus elaphus). Differences in activity synchronization between mixed-sex and unisex groups of red deer explained 35% of the observed degree of intersexual social segregation, showing that activity synchronization is an important cause of social segregation in this species.

Olfaction in the domestic fowl: a critical review.

It has been known for some time that many species of birds, including domestic fowl Gallus domesticus, have an olfactory sense. However, the functional significance of avian olfaction is less clear. We review neurobiological, embryological and behavioral evidence relevant to the question of how domestic fowl use the sense of smell. Evidence suggests a potential role for olfaction in the formation of attachments to familiar objects or environments; in the elicitation of fear responses by alarm and predator-related odors; in the control of feeding and drinking; and in avoidance of noxious substances. The fact that domestic fowl can detect and respond to a wide range of odors, in a variety of behavioral contexts, has important practical implications, especially in relation to welfare and husbandry.

Diversity and substitutability of adjunctive activities under fixed-interval schedules of food reinforcement.

Six rats received food contingent on pressing a lever on fixed-ratio 1, fixed-interval 30-second, and fixed-interval 60-second schedules, with concurrent access to a drinking spout, a running wheel, and a block of wood. Drinking, running, and chewing were monitored automatically, and these and other activities were observed directly during selected sessions. Because all sessions ended after delivery of 60 pellets, total time available for activities other than eating increased over the three schedules. Time spent contacting the lever and visiting the food tray increased in proportion to total available time, whereas the time spent in other activities changed in a complex manner such that drinking was the dominant adjunctive behavior in the 30-second condition, and running or chewing the dominant adjunctive behavior in five of six rats in the 60-second condition. General activity and grooming also occupied significant amounts of time. In a subsequent part of the experiment, running and chewing were prevented, and the majority of other activities, especially drinking and grooming, increased. The results show that (a) FI schedules of food reinforcement are accompanied by a wide variety of adjunctive activities; (b) the preferred activity differs according to the schedule duration; and (c) the extent to which activities substitute for one another is limited by the tendency for different activities to occupy different parts of the interreinforcement interval.

Demand for food on fixed-ratio schedules as a function of the quality of concurrently available reinforcement.

Six rats lever pressed for food on concurrent fixed-ratio schedules, in a two-compartment chamber. In one compartment, mixed diet pellets were delivered on fixed-ratio schedules of 1, 6, 11, and 16; in the other, either no food was delivered, or sucrose or mixed diet pellets were delivered on fixed-ratio 8. The number of pellets obtained in the first compartment declined as a function of fixed-ratio size in that compartment in all three conditions, but the decline was greatest overall with mixed diet pellets concurrently available in the other compartment, and least with no food concurrently available. The result is discussed in terms of economic demand theory, and is consistent with the prediction that elasticity of demand for a commodity (defined in operant terms as the ratio of the proportionate change in number of reinforcements per session to the proportionate change in fixed-ratio size) is greater the more substitutable for that commodity are any concurrently available commodities.

The retained gastric antrum. A case report.

A case of retained gastric antrum, confirmed by radiographic and isotopic techniques, is presented. Any patient with chronic, recurrent peptic ulceration following subtotal gastric resection and gastrojejunostomy should have an adequate upper gastrointestinal series with filling of the afferent loop. In addition, intravenous injection of 99mTc pertechnetate is a noninvasive technique which aids in the identification of a retained gastric antrum.