Optimal group size in territorial animals

Group territoriality is one possible origin of helping behavior in the vertebrates. The fitness of an individual in such a group is modeled as a function of territory quality, unit size, intruder pressure, risk of predation, cost of parental care and sensitivity of territories to resource depletion. All of these variables, as well as social status and sex, are viewed in the perspective of time-energy budgets. Each individual in a population of specified density and distribution of territory qualities is conceived as having a threshold of sociality. Optimal group size is expressed as a compromise between advantages gained by sharing costs and disadvantages arising from resource depletion. The use of time-energy budgets to study group territoriality provides new insights and approaches to the study of helping behavior. The role of resource depletion is singled out by the model for further field work.     

Context-dependent group size choice in fish

The costs and benefits of group membership vary with the size of groups, and individuals are expected to modify their choice of groups in response to ecological factors such as food availability and predation risk. We experimentally examined context-dependent group size choice in a shoaling fish, the banded killifish, Fundulus diaphanus, by using nondirectional odour cues to simulate a food source or a successful attack by a predator (food or alarm treatments) in the laboratory. Group sizes were significantly smaller in the food treatment and larger in the alarm treatment than in control trials. When presented with food and alarm cues together, fish formed groups that were larger than control groups but smaller than those seen with alarm cues alone. These results are consistent with theoretical predictions based on the known benefits and costs of grouping and with previous laboratory work examining the individual shoal choice behaviour of single fish. To examine possible mechanisms of group formation, we developed an individual-based model of shoaling behaviour in which simulated fish were allowed to modify the area over which they interacted with neighbouring individuals. Group size distributions produced by the model were a good approximation of our experimental data. We suggest that local behavioural interaction rules of this type are a potential mechanism by which fish may individually adjust grouping behaviour without requiring extensive information on the position and movement of all possible shoalmates.     

Vigilance and group size in ostriches

Wild ostriches were observed while feeding alone or in groups of up to four birds, and their vigilance (proportion of time with the head up) recorded. Individual vigilance declined as group size increased, mainly through a decrease in the frequency with which the head was raised. Males were more vigilant than females, mainly because they kept their heads up for longer. Vigilance was influenced more by the presence than by the vigilance of a companion. When a head would stay down for a long time was impossible for a predator to predict. It is concluded that grouping by ostriches when feeding results in only a slight reduction in the group's vulnerability to successful predator attack, but in a considerable decrease in individual vulnerability.     

Arbovirus infection increases with group size

Buggy Creek (BCR) virus is an arthropod-borne alphavirus that is naturally transmitted to its vertebrate host the cliff swallow (Petrochelidon pyrrhonota) by an invertebrate vector, namely the cimicid swallow bug (Oeciacus vicarius). We examined how the prevalence of the virus varied with the group size of both its vector and host. The study was conducted in southwestern Nebraska where cliff swallows breed in colonies ranging from one to 3700 nests and the bug populations at a site vary directly with the cliff swallow colony size. The percentage of cliff swallow nests containing bugs infected with BCR virus increased significantly with colony size at a site in the current year and at the site in the previous year. This result could not be explained by differences in the bug sampling methods, date of sampling, sample size of the bugs, age structure of the bugs or the presence of an alternate host, the house sparrow (Passer domesticus). Colony sites that were reused by cliff swallows showed a positive autocorrelation in the percentage of nests with infected bugs between year t and year t+1, but the spatial autocorrelation broke down for year t+2. The increased prevalence of BCR virus at larger cliff swallow colonies probably reflects the larger bug populations there, which are less likely to decline in size and lead to virus extinction. To the authors' knowledge this is the first demonstration of arbovirus infection increasing with group size and one of the few known predictive ecological relationships between an arbovirus and its vectors/hosts. The results have implications for both understanding the fitness consequences of coloniality for cliff swallows and understanding the temporal and spatial variation in arboviral epidemics.     

Optimal group size and northern bobwhite coveys

Northern bobwhite, Colinus virginianus, form social units, called coveys, during the nonbreeding season (approximately September-April). Because the evolutionary advantage of this behaviour is generally unknown, we used controlled group size manipulations within an aviary to investigate whether group size influences (1) the time that the covey spends feeding, (2) the percentage of the covey that is vigilant, (3) the overall vigilance of the group and (4) the time to predator detection. We found that increasing group size increased the time that coveys spent in an exposed feeding area, reduced individual vigilance, improved group vigilance and decreased the time to detection of a potential predator. Additionally, we used experimental reductions of wild northern bobwhite coveys to test whether groups size influences (1) individual and covey survival, (2) daily movement in maintaining covey size and (3) mass change. We conducted field research on 12 independent 259-ha study areas (6 control plots and 6 treatments, where 60% of the population was removed) in east-central Kansas, U.S.A. between 9 November and 31 January, 1997-2000. We radio-marked 386 radiocollared individuals that comprised 137 groups on the study areas. Covey size did not differ between or within years or treatments (X±SE: 10.98±0.22 individuals). Our results indicate that a stable group size existed between 1 and 22 individuals, with 11 being an optimal group size. Small coveys (1-7 individuals) had lower group persistence and individual survival, and used increased movement to create or join larger groups where survival was higher. Large groups (15-22) had lower individual survival, increased group movement and individual mass loss. Density-dependent feedbacks (e.g. lower survival and increased competition) may have lowered larger coveys to a stable size. Our results suggest the regulation of an optimal covey size of 11 was promoted by high group persistence, low group movement, improved feeding efficiency, improved individual predator detection and improved individual survival. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Synergy and group size in microbial cooperation

Abstract Microbes produce many molecules that are important for their growth and development, and the exploitation of these secretions by nonproducers has recently become an important paradigm in microbial social evolution. Although the production of these public-goods molecules has been studied intensely, little is known of how the benefits accrued and the costs incurred depend on the quantity of public-goods molecules produced. We focus here on the relationship between the shape of the benefit curve and cellular density, using a model assuming three types of benefit functions: diminishing, accelerating, and sigmoidal (accelerating and then diminishing). We classify the latter two as being synergistic and argue that sigmoidal curves are common in microbial systems. Synergistic benefit curves interact with group sizes to give very different expected evolutionary dynamics. In particular, we show that whether and to what extent microbes evolve to produce public goods depends strongly on group size. We show that synergy can create an "evolutionary trap" that can stymie the establishment and maintenance of cooperation. By allowing density-dependent regulation of production (quorum sensing), we show how this trap may be avoided. We discuss the implications of our results on experimental design.     

Accuracy and precision of dolphin group size estimates

Estimating the number of dolphins in a group is a challenging task. To assess the accuracy and precision of dolphin group size estimates, observer estimates were compared to counts from large‐format vertical aerial photographs. During 11 research cruises, a total of 2,435 size estimates of 434 groups were made by 59 observers. Observer estimates were modeled as a function of the photo count in a hierarchical Bayesian framework. Accuracy varied widely among observers, and somewhat less widely among dolphin species. Most observers tended to underestimate, and the tendency increased with group size. Groups of 25, 50, 100, and 500 were underestimated by <1%, 16%, 27%, and 47%, respectively, on average. Precision of group size estimates was low, and estimates were highly variable among observers for the same group. Predicted true group size, given an observer estimate, was larger than the observer estimate for groups of more than about 25 dolphins. Predicted group size had low precision, with coefficients of variation ranging from 0.7 to 1.9. Studies which depend on group size estimates will be improved if the tendency to underestimate group size and the high uncertainty of group size estimates are included in the analysis.     

Network cohesion,group size and neocortex size in female-bonded Old World primates

Most primates are intensely social and spend a large amount of time servicing social relationships. In this study, we use social network analysis to examine the relationship between primate group size, total brain size, neocortex ratio and several social network metrics concerned with network cohesion. Using female grooming networks from a number of Old World monkey species, we found that neocortex size was a better predictor of network characteristics than endocranial volumes. We further found that when we controlled for group size, neocortex ratio was negatively correlated with network density, connectivity, relative clan size and proportional clan membership, while there was no effect of neocortex ratio on change in connectivity following the removal of the most central female in the network. Thus, in species with larger neocortex ratios, females generally live in more fragmented networks, belong to smaller grooming clans and are members of relatively fewer clans despite living in a closely bonded group. However, even though groups are more fragmented to begin with among species with larger neocortices, the removal of the most central individual does cause groups to fall apart, suggesting that social complexity may ultimately involve the management of highly fragmented social groups while at the same time maintaining overall social cohesion. These results emphasize a need for more detailed brain data on a wider sample of primate species.     

Adaptive sample size calculations in group sequential trials

A method for group sequential trials that is based on the inverse normal method for combining the results of the separate stages is proposed. Without exaggerating the Type I error rate, this method enables data-driven sample size reassessments during the course of the study. It uses the stopping boundaries of the classical group sequential tests. Furthermore, exact test procedures may be derived for a wide range of applications. The procedure is compared with the classical designs in terms of power and expected sample size.     

Social group size predicts the evolution of individuality

Discriminating among individuals is a critical social behavior in humans and many other animals and is often required for offspring and mate recognition, territorial or coalitional behaviors, signaler reliability assessment, and social hierarchies. Being individually discriminated is more difficult in larger groups, and large group size may select for increased individuality-signature information-in social signals, to facilitate discrimination. Small-scale studies suggest that more social species have greater individuality in their social signals, such as contact calls. However, this relationship has not been evaluated in a broader-scale evolutionary context or in social signals other than contact calls. It is not yet known whether social group size may be viewed as a general evolutionary driver of individuality. Here we show a strong positive evolutionary link between social group size in sciurid rodents and individuality in their social alarm calls. Social group size explained over 88% of the variation in vocal individuality in phylogenetic independent contrasts. Species living in larger groups, but not in more complex groups, had more signature information in their calls. Our results suggest that social group size may promote the evolution of individual signatures and that the sociality-individuality relationship may be a general phenomenon in nature.     

Policing effectiveness depends on relatedness and group size

Cohesion of social groups requires the suppression of individual selfishness. Indeed, worker egg laying in insect societies is usually suppressed or punished through aggression and egg removal. The effectiveness of such "policing" is expected to increase with decreasing relatedness, as inclusive fitness of group members is more strongly affected by selfish worker reproduction when group members are less closely related to each other. As inclusive fitness is also influenced by the costs and benefits of helping, the effectiveness of policing should decrease with increasing colony size, because the costs for the whole colony from selfish worker reproduction are proportionally reduced in large groups. Here, we show that policing effectiveness in colonies of the ant Temnothorax unifasciatus is low in large groups and high in small groups when relatedness is high. When we experimentally decreased the relatedness in groups, the policing effectiveness reached the same high level as in small, highly related groups, irrespective of group size. Therefore, our results indicate that policing effectiveness is simultaneously shaped by relatedness and group size, that is, an ecological factor. This may have major implications for testing policing across species of animals.     

Short communication: rat's demand for group size

Social isolation compromises the welfare of rats. However, it is not clear how many rats should be housed together under laboratory conditions. Pair housing, sometimes recommended over group housing, may help avoid aggression and disease transmission. Female rats, however, showed the highest average demand for a group size of 6 (versus 1, 2, 4, and 12) when stocking density was maintained at 20 cm2/rat. This finding contributes to work suggesting that rats should be group housed. This article shows that further studies are required into the actual risks of disease and injury associated with group versus pair housing.     

To group or not to group: group size dynamics and intestinal parasites in Indian peafowl populations

acta ethologica - Animals can form groups for various reasons including safety from predators, access to potential mates and benefits of allo-parental care. However, there are costs associated with...     

Modification of sample size in group sequential clinical trials

In group sequential clinical trials, sample size reestimation can be a complicated issue when it allows for change of sample size to be influenced by an observed sample path. Our simulation studies show that increasing sample size based on an interim estimate of the treatment difference can substantially inflate the probability of type I error in most practical situations. A new group sequential test procedure is developed by modifying the weights used in the traditional repeated significance two-sample mean test. The new test has the type I error probability preserved at the target level and can provide a substantial gain in power with the increase of sample size. Generalization of the new procedure is discussed.     

Squirrel monkey dominance and social behavior as related to group size and group structure

Adult male and female squirrel monkeys were tested in nonsocial adaptation and pairwise and triad social situations differing in sex composition. Social behaviors, nonsocial behaviors, and dominance hierarchies were observed during social testing. Dominance hierarchies were similar in groups differing in size and social structure. Nonsocial behaviors decreased in females and submissive animals paired with males or dominant monkeys. Aggressiveness between females decreased and the beginnings of coalitions between females were observed in the presence of a male. The social behavior patterns, but not dominance hierarchies, are consistent with behaviors observed in larger groups of squirrel monkeys.