Geometry for mutualistic and selfish herds: the limited domain of danger

We present a two-dimensional individual-based model of aggregation behaviour in animals by introducing the concept of a "limited domain of danger", which represents either a limited detection range or a limited attack range of predators. The limited domain of danger provides a suitable framework for the analysis of individual movement rules under real-life conditions because it takes into account the predator's prey detection and capture abilities. For the first time, a single geometrical construct can be used to analyse the predation risk of both peripheral and central individuals in a group. Furthermore, our model provides a conceptual framework that can be equally applied to aggregation behaviour and refuge use and thus presents a conceptual advance on current theory that treats these antipredator behaviours separately. An analysis of individual movement rules using limited domains of danger showed that the time minimization strategy outcompetes the nearest neighbour strategy proposed by Hamilton's (J. Theor. Biol. 31 (1971) 295) selfish herd model, whereas a random strategy confers no benefit and can even be disadvantageous. The superior performance of the time minimization strategy highlights the importance of taking biological constraints, such as an animal's orientation relative to its neighbours, into account when searching for efficient movement rules underlying the aggregation process.     

Visual sensitivity and foraging in social wasps

Summary While there is a distinction between that intensity of illumination which permits social wasps to forage, and that to which a sessile worker can respond, nevertheless illumination is the most critical of the environmental factors which control the activity of wasps. Low temperatures, high winds, and heavy rain all reduce activity but unless exceptionally severe do not wholly stop it. At dawn, when the critical level of illumination is attained, workers leave the nest, but at dusk they will not leave should the same critical level be due in the course of the foraging flight, after which they could not return.The three species of wasp,Vespula vulgaris, V. rufa, andV. germanica have a common threshold of illumination, although the hornet,Vespa crabro can forage in moonlight at an altogether lower illumination. Honey-bees normally need a still higher illumination than do wasps.In all these species, the thresholds of illumination are related to the length of the compound eyes, so that species with large eyes need less light by which to forage. Moreover, there is a slight difference between the threshold at dawn when workers leave the nest, and that at dusk, when they must needs have sufficient light by which to return. This difference is almost constant for each species, when, as is customary, one measures it on a logarithmic scale.Lastly, the estimates, which these experiments provide, of the threshold illuminations depend stochastically on the number of workers foraging. A correction for this bias is given.

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Sommaire Parmi les facteurs du milieu qui contrôlent l'activité des guêpes, celui de l'intensité de lumière est le plus important; toutefois on note une différence entre l'intensité de lumière qui permet aux guêpes sociales de fourrager, et celle qui produit une réponse des ouvrières sessiles.En général, les basses températures, les vents forts, et les grandes pluies réduisent leur activité, mais ces facteurs ne l'arrêtent pas complètement, à moins qu'ils ne soient très marqués.A l'aube, quand le niveau critique de lumière est atteint, les ouvrières quittent le guêpier, mais, le soir, si elles s'attendent à ce que la lumière vienne à s'abaisser au cours de leur sortie au-dessous du niveau critique, elles ne sortent pas.Les trois espèces de guêpe,Vespula vulgaris, V. rufa, etV. germanica, réagissent au même seuil de lumière, mais le frelon,Vespa crabro, est capable de fourrager au clair de lune par une lumière moins intense. Normalement, les abeilles exigent une lumière plus intense que les guêpes.Dans toutes ces espèces, le seuil de lumière se rapporte à la hauteur des yeux composés, par conséquent les espèces pourvues de grands yeux sont à même de fourrager par une lumière moins intense. De plus, il y a une légère différence entre le seuil de lumière à l'aube, quand les ouvrières quittent le guêpier, et celui du soir lorsqu'elles ont besoin d'une lumière suffisante pour rentrer. Cette différence, quand elle est mesurée à l'échelle logarithmique, comme il est d'usage, est presque constante pour chaque espèce.Enfin, les évaluations du seuil de lumière dans ces expériences dépendent stochastiquement du nombre d'ouvrières en train de fourrager. On a tenu compte de ce fait.

     

Visual perception and social foraging in birds

Birds gather information about their environment mainly through vision by scanning their surroundings. Many prevalent models of social foraging assume that foraging and scanning are mutually exclusive. Although this assumption is valid for birds with narrow visual fields, these models have also been applied to species with wide fields. In fact, available models do not make precise predictions for birds with large visual fields, in which the head-up, head-down dichotomy is not accurate and, moreover, do not consider the effects of detection distance and limited attention. Studies of how different types of visual information are acquired as a function of body posture and of how information flows within flocks offer new insights into the costs and benefits of living in groups.     

Brain's neural switch for social dominance in animals

正Animals perform social behaviors during their lives to survive and reproduce. As one of the most robust and fundamental social behaviors, social dominance determines individual's behavioral displays, priority access to food, mate,territory or other resources, and impacts on its physical and mental health. In human society, socioeconomic status (SES)is a major predictor of physical and mental outcomes, even when the participants have equal access to health care. A Norwegian scientist Thorleif Schjelderup-Ebbe first scientifically described dominance behaviors when he observed     

Departure rules used by bees foraging for nectar: A field test

Summary Departure rules used by solitary long-tongued bees (Anthophora spp. andEucera spp.) collecting nectar from flowers ofAnchusa strigosa (Boraginaceae) were studied. The amount of nectar a bee receives from an individual flower was estimated by measuring the time elapsed since the previous bee visit to that flower. Measurements of nectar accumulation in experimentally emptied flowers indicated that this time interval is an accurate predictor of nectar volumes in flowers. We found that nectar rewards influence the probability of departure from individual plants, as well as distances of movements within plants. The probability of departure from individual plants was negatively related to the amount of reward received at the two lastvisited flowers. This result indicates that the bees used a probabllistic departure rule, rather than a simple threshold departure rule, and that rewards from both the current and the previously visited flower were important in determining departure points. Distances of inter-flower movements within plants were negatively related to the amount of reward received at the current flower. The overall results suggest that the pollinators ofA. strigosa make two types of departure decisions-departures from the whole plant and departures from the neighbourhood of individual flowers-and that they use different departure rules for each scale. Factors influencing the decision-making processes of the observed foraging behaviour are discussed.     

The advantages of social foraging in downy woodpeckers

I investigated the advantages gained by downy woodpeckers (Picoides pubescens) which join mixed-species winter flocks. Woodpeckers foraging alone showed high levels of vigilance as measured by head-cocking rates, and low feeding rates. Woodpeckers foraging with one or two flock members showed intermediate rates of head-cocking and feeding, while woodpeckers foraging with flocks of three or more birds showed low head-cocking rates and high feeding rates. Although local enhancement and copying may contribute to the woodpeckers' increased foraging efficiency in a flock, these do not appear to be the main factors. As downy woodpeckers spend less time on vigilance, they devote more time to foraging, thereby increasing their foraging efficiency     

Flexible search tactics and efficient foraging in saltatory searching animals

Summary Foraging is one of the most important endeavors undertaken by animals, and it has been studied intensively from both mechanistic-empirical and optimal foraging perspectives. Planktivorous fish make excellent study organisms for foraging studies because they feed frequently and in a relatively simple environment. Most optimal foraging studies of planktivorous fish have focused, either on diet choice or habitat selection and have assumed that these animals used a cruise search foraging strategy. We have recently recognized that white crappie do not use a cruise search strategy (swimming continuously and searching constantly) while foraging on zooplankton but move in a stop and go pattern, searching only while paused. We have termed thissaltatory search. Many other animals move in a stop and go pattern while foraging, but none have been shown to search only while paused. Not only do white crappie search in a saltatory manner but the components of the search cycle change when feeding on prey of different size. When feeding on large prey these fish move further and faster after an unsuccessful search than when feeding on small prey. The fish also pause for a shorter period to search when feeding on large prey. To evaluate the efficiency of these alterations in the search cycle, a net energy gain simulation model was developed. The model computes the likelihood of locating 1 or 2 different size classes of zooplankton prey as a function of the volume of water scanned. The volume of new water searched is dependent upon the dimensions of the search volume and the length of the run. Energy costs for each component of the search cycle, and energy gained from the different sized prey, were assessed. The model predicts that short runs produce maximum net energy gains when crappie feed on small prey but predicts net energy gains will be maximized with longer runs when crappie feed on large prey or a mixed assemblage of large and small prey. There is an optimal run length due to high energy costs of unsuccessful search when runs are short and reveal little new water, and high energy costs of long runs when runs are lengthy. The model predicts that if the greater search times observed when crappie feed on small prey are assessed when they feed on a mixed diet of small and large prey, net energy gained is less than if small prey are deleted from the diet. We believe the model has considerable generality. Many animals are observed to move in a saltatory manner while foraging and some are thought to search only while stationary. Some birds and lizards are, known to modify the search cycle in a manner similar to white crappie.Components of the search cycle and dimensions of the location space SST (sec) Successful search time — the average time stationary prior to a pursuit - USST (sec) Unsuccessful search time — the average time stationary prior to a run - PT (sec) Pursuit time-PL/SS — the time to pursue prey at a given distance away. It is calculated by dividing the pursuit distance by swim speed - RT (sec) Run time-RL/SS — the time to complete a run of a given length. It is calculated by dividing the run length, by swim speed - PL (cm) Pursuit length-distance moved to attack prey - RL (cm) Run length-distance moved between consecutive searches - SS (cm/sec) Swim speed — the speed of movement during a pursuit or run - LS (l) Location space — the area or volume within which prey are located. In the case of white crappie the search space is shaped like a pie wedge with the fish positioned at the apex of the wedge - LA (^o) Location angle—the angle of the wedge-shaped search space - LH (cm) Location height—the height of the wedge-shaped search space - LD (cm) Location distance—the length of long axis of the wedge-shaped search space. Components of the location probability model RND Random number-random number generated through BASICA - SV (l) Search volume—the volume of water actually searched after one run of given length - SV_MAX (l) Maximum search volume—the greatest search volume that can be based upon LA, LH, LD and unaffected by the previous search - SV_R (l) Search volume researched—that volume of SV_MAX that is researched where RLo Search volume unsearched—that volume of SV_MAX not previously searched - AD (#/1) Absolute density—the density of zooplankton prey in numbers per liter - VD (#) Visual density—the number of zooplankton prey in the search volume - LP (%) Location probability—the probability that one or more prey are in the search volume Components of the net energy gain model NEG (cal/sec) Net energy gain-total calories ingested, less total calories used, divided by total time. - E _e (cal) Energy expended on the search cycle - E _i (cal) Energy intake - e _p (cal) Energy content of a given individual prey - P _i Total number of prey ingested - e _r (cal) Energy expended while searching - e _s (cal) Energy expended while swimming - T _t (sec) Total time-time expended to eat a given number of prey     

A selfish origin for recombination

Recent findings of molecular biology show that recombination is initiated by interactions between homologous chromosomes and that an allele can induce the initiation of recombination on the homolog. Since gene conversion at the site of initiation is strong enough to promote the transmission of that allele, recombination may be a way for a self-promoting element to spread, even if it gives no advantage to the individual or to the population. I develop a simple model and discuss available molecular evidence in support of this hypothesis. A consequent argument is that with asexual reproduction the evolution of recombination leads to an intragenomic conflict, and a possible outcome of this conflict may be the origin of sexual reproduction.     

Co-evolution of learning complexity and social foraging strategies

Variation in learning abilities within populations suggests that complex learning may not necessarily be more adaptive than simple learning. Yet, the high cost of complex learning cannot fully explain this variation without some understanding of why complex learning is too costly for some individuals but not for others. Here we propose that different social foraging strategies can favor different learning strategies (that learn the environment with high or low resolution), thereby maintaining variable learning abilities within populations. Using a genetic algorithm in an agent-based evolutionary simulation of a social foraging game (the producer-scrounger game) we demonstrate how an association evolves between a strategy based on independent search for food (playing a producer) and a complex (high resolution) learning rule, while a strategy that combines independent search and following others (playing a scrounger) evolves an association with a simple (low resolution) learning rule. The reason for these associations is that for complex learning to have an advantage, a large number of learning steps, normally not achieved by scroungers, are necessary. These results offer a general explanation for persistent variation in cognitive abilities that is based on co-evolution of learning rules and social foraging strategies.     

Ontogenetic shifts within the selfish herd: predation risk and foraging trade-offs change with age in colonial web-building spiders

We tested for the existence of density dependence in annual adult rangeland grasshopper (Orthoptera: Acrididae) data from Montana, USA (1951–1991). Statistical density dependence was, in the sense of a stochastic equilibrium or return tendency, detected in all of the grasshopper mean density time-series from the three major physiographic regions of the state, Northern Glaciated Plains, Southern Unglaciated Plains, and Western Mountains. Parameters were estimated for a model that described the stochastic equilibrium characteristics of regional mean densities. The analyses showed that rangeland grasshopper regional densities fluctuate according to gamma distribution with a mean of 6.1–6.3 grasshoppers per m². Further, when regions exhibit outbreaks, the resulting infestation period (duration of outbreak) is short, spanning only a few generations.     

The social cognition of social foraging: partner selection by underlying valuation

Humans and other animals have a variety of psychological abilities tailored to the demands of asocial foraging, that is, foraging without coordination or competition with other conspecifics. Human foraging, however, also includes a unique element: the creation of resource pooling systems. In this type of social foraging, people contribute when they have excess resources and receive provisioning when in need. Is this behavior produced by the same psychology as asocial foraging? If so, foraging partners should be judged by the same criteria used to judge asocial patches of resources: the net energetic benefits they provide. The logic of resource pooling speaks against this. Maintaining such a system requires the ability to judge others not on their short-term returns, but on the psychological variables that guide their behavior over the long term. We test this idea in a series of five studies using an implicit measure of categorization. Results showed that (a) others are judged by the costs they incur (a variable not relevant to asocial foraging), whereas (b) others are not judged by the benefits they provide when benefits provided are unrevealing of underlying psychological variables (despite this variable being relevant to asocial foraging). These results are suggestive of a complex psychology designed for both social and asocial foraging.     

Exploring foraging decisions in a social primate using discrete-choice models

Abstract There is a growing appreciation of the multiple social and nonsocial factors influencing the foraging behavior of social animals but little understanding of how these factors depend on habitat characteristics or individual traits. This partly reflects the difficulties inherent in using conventional statistical techniques to analyze multifactor, multicontext foraging decisions. Discrete-choice models provide a way to do so, and we demonstrate this by using them to investigate patch preference in a wild population of social foragers (chacma baboons Papio ursinus). Data were collected from 29 adults across two social groups, encompassing 683 foraging decisions over a 6-month period and the results interpreted using an information-theoretic approach. Baboon foraging decisions were influenced by multiple nonsocial and social factors and were often contingent on the characteristics of the habitat or individual. Differences in decision making between habitats were consistent with changes in interference-competition costs but not with changes in social-foraging benefits. Individual differences in decision making were suggestive of a trade-off between dominance rank and social capital. Our findings emphasize that taking a multifactor, multicontext approach is important to fully understand animal decision making. We also demonstrate how discrete-choice models can be used to achieve this.     

An invitation to die: initiators of sociality in a social amoeba become selfish spores

Greater size and strength are common attributes of contest winners. Even in social insects with high cooperation, the right to reproduce falls to the well-fed queens rather than to poorly fed workers. In Dictyostelium discoideum, formerly solitary amoebae aggregate when faced with starvation, and some cells die to form a stalk which others ride up to reach a better location to sporulate. The first cells to starve have lower energy reserves than those that starve later, and previous studies have shown that the better-fed cells in a mix tend to form disproportionately more reproductive spores. Therefore, one might expect that the first cells to starve and initiate the social stage should act altruistically and form disproportionately more of the sterile stalk, thereby enticing other better-fed cells into joining the aggregate. This would resemble caste determination in social insects, where altruistic workers are typically fed less than reproductive queens. However, we show that the opposite result holds: the first cells to starve become reproductive spores, presumably by gearing up for competition and outcompeting late starvers to become prespore first. These findings pose the interesting question of why others would join selfish organizers.     

Beyond learning fixed rules and social cues: abstraction in the social arena

Abstraction is a central idea in many areas of physical comparative cognition such as categorization, numerical competence or problem solving. This idea, however, has rarely been applied to comparative social cognition. In this paper, I propose that the notion of abstraction can be applied to the social arena and become an important tool to investigate the social cognition and behaviour processes in animals. To make this point, I present recent evidence showing that chimpanzees know about what others can see and about what others intend. These data do not fit either low-level mechanisms based on stimulus-response associations or high-level explanations based on metarepresentational mechanisms such as false belief attribution. Instead, I argue that social abstraction, in particular the development of concepts such as seeing in others, is key to explaining the behaviour of our closest relative in a variety of situations.     

Advantages of social foraging of Willow Tits Parus montanus

The mean number of Willow Tits Parus montanus in single-species flocks was significantly larger than in mixed-species flocks of Willow and Coal Tits P. ater. Both flock size and the tendency of Willow Tits to join mixed-species flocks were negatively correlated with ambient temperature, probably because each bird, when the metabolic rate of the birds increased, could allocate more time to foraging due to improved predator detection by many eyes. The vigilance time of Willow Tits decreased with flock size and was determined by the total number of individuals in a flock rather than by the number of Willow Tits in mixed-species flocks of Willow and Coal Tits.     

Prepared social learning about dangerous animals in children

Natural selection is likely to have shaped developmental systems for rapid acquisition of knowledge about environmental dangers, including dangerous animals. However, learning about dangerous animals through direct encounters can be costly and potentially fatal. In social species such as humans, the presence of stored information about danger in the minds of conspecifics might favor the evolution of prepared social learning mechanisms that cause children to preferentially attend to and remember culturally transmitted information about danger. Here we use an experimental learning task to show that children from two very different cultures exhibit prepared social learning about dangerous animals: city-dwelling children from Los Angeles, who face relatively little danger from animals, and Shuar children from the Amazon region of Ecuador, to whom dangerous animals pose a much greater threat. Both populations exhibited similar prepared learning effects. Danger information was learned in a single trial without feedback, immediately entered long-term memory, and was recalled with only minor attenuation a week later, while other information presented at the same time (animal names and diets) was immediately forgotten. We discuss the significance of these design features of prepared learning in light of the phylogeny and function of danger learning systems.