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.      

Optimal group size and seasonal stress in ring-tailed lemurs (Lemur catta)

Adaptive explanations for social grouping assume that thereare fitness consequences associated with group size, and individualsmaintain membership in groups of favorable size to maximizefitness. Here I examine fecal cortisol concentrations as a hormonalmeasure of stress to assess the relative well-being of Lemurcatta in groups of different size and in seasons of normal andlow tamarind fruit availability. I test the hypotheses thatthere is an optimal group size at which cortisol is lowest andthat optimal group size changes in food-scarce conditions. Icollected 799 fecal samples from 87 individuals in seven free-rangingL. catta groups at Berenty Reserve, Madagascar, over a 1-yearperiod (August 1999–July 2000) and determined fecal cortisolconcentrations using a radioimmunoassay. Expressing these asresiduals from monthly population means to control for temporalfluctuations in cortisol concentration, I calculated mean fecalcortisol levels for each animal in seasons of normal and lowtamarind fruit abundance and over the entire year. Overall,females exhibited lowest mean cortisol levels in groups of intermediatesize, suggesting that there are benefits to maintaining membershipin these groups. Females in groups that were atypically largeor small for their habitat type had higher mean cortisol levelsthan typical groups. Cortisol levels increased in food-scarceconditions for larger groups, suggesting that intergroup competitiveadvantages do not outweigh intragroup feeding competition atthis time. Group size may be optimized for long-term averageconditions, and short-term stresses may intermittently alterthe costs associated with group size.     

群居动物中的共同决策

群居的个体能够更好的躲避天敌的捕食,增强与同种其它种群的资源竞争,增加生殖机会与生殖成功率,使信息在个体之间顺畅的交流等。但是群内个体又会在有限的资源下产生群内的个体竞争,从而引起动物群的分裂,失去群居的利益。为了保持动物群的聚合力,当动物群内出现冲突时,需要在群内个体间形成协调各种冲突以及在群内达成共识的机制。依据动物社群组成的不同,领导力机制和自组织机制是协调群内冲突,形成一致决策和联合决策的两种主要机制。从行为生态学的角度入手,阐述了动物群内共同决策的类型,群内冲突的协调机制,共同决策的制定过程,比较了信息因素与利益因素在共同决策制定过程中的影响。旨在藉此推进我国相关领域研究的开展, 为群居动物社会与行为生态学研究者启发思路,并为保护群居濒危物种提供科学依据。     

Persistence of disease in territorial animals: insights from spatial models of Tb

Early models of directly transmitted wildlife disease focused on rabies transmission as a travelling wave, usually in a homogeneous density of wildlife. Such models of epi-enzootic diseases paid little attention to local-scale disease prevalence. Historical data on bovine tuberculosis (Tb) in cattle indicates that very localised areas can suffer from frequent repeat breakdowns, indicating that some environmental factors might be the cause. There are a number of different ways to simulate such local disease ‘hotspots’ in wildlife, and these resultant hotspots may mean that, overall, wildlife disease prevalence is very low. However, spatial and temporal persistence of this hotspot is more difficult to model. This heterogeneity in disease prevalence is difficult to produce in non-spatial models, and is one of the reasons why such models gave poor predictions of disease dynamics in the field. For example, Nigel Barlow struggled with finding a way to produce this spatial heterogeneity in mathematical models, culminating in his 2000 paper in Journal of Animal Ecology. This gave a phenomenological treatment, but not a causative solution. I take a look at the various causative methods of producing disease heterogeneity in simulation models of Tb, a chronic wildlife disease. These include (1) chance, (2) model artefacts, (3) population (e.g. demographic, genetic) heterogeneity and (4) environmental heterogeneity. I further argue that only (4) can be predicted over a medium timescale, and propose methods to assess the contribution of (1) and (2) in a model. I also discuss how spatial heterogeneity may affect Tb management.     

Optimal body size in bumblebees

Summary It is hypothesized that the body size of a bumblebee will be that size which maximizes its average net rate of energy intake while collecting nectar. A mathematical model is developed with the result that the net rate of energy intake of a nectar-collecting bumblebee is expressed as a function of the body size of the bumblebee. From this model the body size which maximizes the net rate of energy intake (i.e., optimal body size) is found (as the solution of an implicit equation). In this situation the advantage of large size is that larger bumblebees fly faster and hence take less flight time than smaller bumblebees. The disadvantage of larger size is greater energetic costs.The parameters of the model are estimated using data obtained from the foraging behavior of bumblebees on monkshood (Aconitum columbianum). The optimal body size is then calculated for workers of Bombus appositus which obtained almost all their nectar from monkshood. The observed and expected (i.e., optimal) body size are found to be close and not significantly different.The model also predicts that, from the bumblebee's point of view, there should be a positive correlation between the size of the bumblebee and the average amount of nectar obtained per flower. Evidence of this correlation is presented and the possible significance of the correlation from the plant's point of view is discussed. A possible extension of the model to general relationships between predator body size, prey size and prey density is discussed.     

Body patterns as potential amplifiers of size and condition in a territorial spider

Signal costs, and associated trade-offs, are widely thought of as fundamental to the evolution and maintenance of reliable signals. Contrasting this view, the amplifier mechanism postulates that signals may be deemed reliable by design rather than by costs. Although the idea is not new, investigation of amplifier signals in nature remains scant. Here we show that displays and body markings on the Mediterranean tarantula, Lycosa tarantula (L.) (Lycosidae), are consistent with interpretation as amplifiers of size and body condition. The size of tibia and abdomen ventral markings (which alternate between dark and pale patterns) presented to rivals during contests are highly correlated with body size and condition, respectively. Abdomen markings have the additional feature of placement on a region especially influenced by abdomen distension and therefore has greater proportional variation than the underlying amplified trait. In spite of the differences in size and in body patterns between spiders of two isolated and ecologically different populations, correlation between body marking and dimensions, and relatively high proportional variation of the abdomen markings, was maintained, probably reflecting the adaptive value of these traits. We conducted a feeding experiment which showed that the abdominal markings enlarged (stretched with abdomen distension) at a higher rate if spiders were provided with more food, further supporting the idea that this marking amplifies body condition.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 355–364.     

Sexual selection and sexual size dimorphism in animals

Sexual selection is often considered as a critical evolutionary force promoting sexual size dimorphism (SSD) in animals. However, empirical evidence for a positive relationship between sexual selection on males and male-biased SSD received mixed support depending on the studied taxonomic group and on the method used to quantify sexual selection. Here, we present a meta-analytic approach accounting for phylogenetic non-independence to test how standardized metrics of the opportunity and strength of pre-copulatory sexual selection relate to SSD across a broad range of animal taxa comprising up to 95 effect sizes from 59 species. We found that SSD based on length measurements was correlated with the sex difference in the opportunity for sexual selection but showed a weak and statistically non-significant relationship with the sex difference in the Bateman gradient. These findings suggest that pre-copulatory sexual selection plays a limited role for the evolution of SSD in a broad phylogenetic context.     

Behavioral ecology of disturbed landscapes: the response of territorial animals to relocation

Intraspecific territorial interactions are common for a largevariety of wildlife species. This often results in high-qualityhabitat being occupied by dominant individuals, with subordinatesrelegated to lower quality habitat. The role that these territorialinteractions play in influencing the redistribution of animalsthat have been evicted from their native home ranges remainsunclear. My goals were to determine (1) how the density of conspecificsin the new habitats impacts resettlement patterns and (2) towhat extent prior dominance status is maintained when an animalis forced to relocate. I relocated white-footed mice (Peromyscusleucopus) from high-quality, oak-dominated hardwood and lowerquality, white pine forests to novel sites and released themalong the ecotone of these two habitat types. I relocated micefirst in the presence and then in the absence of a natural densityof resident mice. Habitat selection and resource acquisitionof relocated mice were assessed via mark-recapture livetrappingand passive integrated transponder tagging. Relocated mice selectedhigh-quality habitat significantly more often when residentmice were absent, illustrating the importance of territorialinteractions for determining resettlement patterns of relocatedindividuals. Data on resource acquisition also reflected thecompetitive influence of resident mice—relocated micewere significantly more successful accessing food resourcesin the treatment without residents. The habitat of origin didnot significantly impact habitat selection or resource acquisition,indicating that all relocated individuals were at a disadvantagecompared to residents.     

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.     

How territorial animals compete for divisible space: a learning-based model with unequal competitors

It is widely assumed that aggressive behavior affects space acquisition in territorial species, but to date most workers have focused on competition for indivisible space, that is, space that cannot be divided or shared. We present a learning-based model that investigates the effects of aggressive interactions on space acquisition when unequal competitors arrive and settle in patches of divisible space. This model assumes that aggressive interactions act as punishment, in the sense that previous aggressive interactions in a given area reduce an individual's likelihood of returning to that area. Individually based, spatially explicit simulations incorporating this and other assumptions were used to investigate the effects of different types of aggressive interactions on the space use of individuals and dyads settling in divisible space. At the individual level, final space use was related to the amount of punishment that individuals inflicted on their opponents during aggressive interactions; in general, highly aggressive individuals acquired larger, more exclusive home ranges than less aggressive individuals. At the dyadic level, the division or sharing of space between competitors depended on both the relative and absolute punishment that competitors inflicted on one another during aggressive interactions. Aggressive interactions in which both participants strongly punished one another (e.g., escalated fights) produced mutually exclusive home ranges, interactions with intermediate levels of punishment produced asymmetrical space use patterns proportional to asymmetries in punishment levels, and interactions involving little punishment for either participant generated large home ranges with extensive home range overlap. Overall, our model implies that territorial animals need not "win" aggressive interactions to win divisible space, that repeatedly "nagging" an opponent may also be a viable strategy for gaining space, and that a learning-based approach can account for puzzling patterns in the territorial literature, for example, observations of individuals who acquire space by initiating aggressive interactions that they never win.     

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.     

The relationship between body size and population abundance in animals

On average, large-bodied species live at lower densities than small-bodied ones. Early studies suggested that population densities might scale so that the energy use of a population is independent of body size. However, recent work shows that, at the scale of local communities, this is rarely true and that the pattern varies among taxonomic or ecological subsets of those communities. Energetic considerations may only be relevant to the densities of more abundant species. In fact, within natural assemblages o f organisms, the underlying relationship is very variable; in subsets of those assemblages, ecological processes such as competition may structure abundance patterns.     

Male size does not affect territorial behaviour and life history traits in a sphecid wasp

The optimum maternal investment per offspring is determined by the relationship between the investment per offspring and offspring fitness. In the European beewolf, Philanthus triangulum, a solitary mass-provisioning sphecid wasp, offspring size correlates with the amount of provisions. We investigated whether the reproductive success of adult males depends on body size in a way that would influence the allocation of parental investment. Since the mating success of P. triangulum males cannot be determined by observation in the field, we assessed the influence of male size on characteristics of their territories, territorial behaviour and life history traits. Territory size was weakly correlated with male size, but a measure of territory quality (number of female nests in the vicinity) was independent of male size. Neither the duration of ownership nor the intensity of scent marking was correlated with male size. Territory owners were slightly smaller than nonterritorial males. The absolute amount of fat was positively correlated with size but, owing to allometric relationships, the energetic equivalent of the fat store appeared to be independent of size. Life span was not significantly influenced by body size under four different conditions (with and without food in the laboratory, in an outdoor flight cage and in the field). We discuss the discrepancy between these results and other studies that have mostly reported advantages to large males. We suggest that in noncontact male-male interactions, as seen in the European beewolf, body size might not be the key determinant for success in contests. We conclude that there is no evidence for a strong size dependence of male reproductive success. Thus the reproductive success of male progeny probably does not depend on parental investment in a way that would influence the investment allocation of females. Copyright 2000 The Association for the Study of Animal Behaviour.     

Optimal hunting group size: the need for lions to defend their kills against loss to spotted hyaenas

This paper presents preliminary evidence that kleptoparasitism by spotted hyaenas may influence optimal hunting group size of lions. In the absence of adult male lions, hyaenas can drive female and subadult lions off their kills provided they outnumber the lions by a factor of four. Hence the larger the group of lions on the kill, the greater their chance of defending their food against invading hyaenas. At Savuti, where there was a shortage of adult male lions, the groups of female and subadult lions lost almost 20% of their food to hyaenas. Losses were most frequent for those living in small groups. These lions were often satiated by the time the hyaenas acquired the kill, so the hyaenas did not cause immediate need, yet constituted a constant energy drain on lions by forcing them to hunt more frequently. The implications of these observations for modelling optimal hunting group size of lions are discussed.