Patch exploitation, group foraging, and unequal competitors

Charnov's (1976) marginal value theorem, MVT, addresses howlong a forager should stay in a patch of prey to maximize itsgain. Information-sharing models of group foraging suggest thatindividuals should join groups to improve their patch-findingrate. This is achievable if group members share informationabout the location of food patches. The determinants of theMVT are searching time and cumulative gain against time in apatch, those of the group foraging models are searching time,group size, and individual differences in ability to monopolizethe prey found. After combining the MVT and information-sharingmodels we explore the consequences of unequal competitors (good,G, and poor, P) foraging in groups. Under this domain G andP differ in their accumulated harvest against time in a patch.When the gain function of P is obtained by mere scaling of thatof G, optimal patch residence times for individuals of the twophenotypes do not differ. However, if the gain functions ofG and P cannot be derived from each other by a constant scalingmultiplier, the optimal patch times for G and P are not necessarilythe same. Under these conditions the model suggests that foraginggroups should become assorted by foraging ability.     

Refuge sites of voles under owl predation risk: priority of dominant individuals?

In an aviary experiment, we studied whether body size or habitatfamiliarity of field voles (Microtus agrestis) affected predationrisk by Tengmalm's owls (Aegolius funereus). In the field, wecompared the body size of field voles snap-trapped in good (covered)and poor (open) habitats in 1992 and 1994 to determine whetherthere were habitat-related differences in the body size of voles.In the aviary, large individuals occupied the good habitat significantlymore than small individuals both in the control (owl not present)and experimental treatments (owl present). Furthermore, habitat-familiarvoles inhabited the good habitat more than habitat-unfamiliarvoles did when an owl was present Our field data were consistentwith our aviary data: larger field voles were more frequentlyfound in good habitats than in poor habitats. In the aviary,Tengmalm's owl predation risk was higher for small and habitat-unfamiliarvoles. This suggests that large field voles may have priorityto sheltered habitats. Furthermore, habitat familiarity mayplay a central role in avoiding risky habitats.     

Ecological and social determinants of group size in transient killer whales

Most analyses of the relationship between group size and foodintake of social carnivores have shown a discrepancy betweenthe group size that maximizes energy intake and that which ismost frequently observed. Around southern Vancouver Island,British Columbia, killer whales of the so-called transient formforage in small groups, and appear to prey exclusively on marinemammals. Between 1986 and 1993, in approximately 434 h of observationson transient killer whales, we observed 138 attacks on fivespecies of marine mammals. Harbor seals were most frequentlyattacked (130 occasions), and the observed average energy intakerate was more than sufficient for the whale's energetic needs.Energy intake varied with group size, with groups of three havingthe highest energy intake rate per individual. While groupsof three were most frequently encountered, the group size experiencedby an average individual in the population (i.e., typical groupsize) is larger than three. However, comparisons between observedand expected group sizes should utilize only groups engagedin the behavior of interest. The typical size of groups consistingonly of adult and subadult whales that were engaged primarilyin foraging activities confirms that these individuals are foundin groups that are consistent with the maximization of energyintake hypothesis. Larger groups may form for (1) the occasionalhunting of prey other than harbor seals, for which the optimalforaging group size is probably larger than three; and (2) theprotection of calves and other social functions. Key words:dispersal, foraging, group hunting, harbor seals, killer whales,optimal group size, social structure. [Behav Ecol 7: 408-416(1996)]     

Advantages of social foraging in crab spiders: Groups capture more and larger prey despite the absence of a web

Among group‐living spiders, subsocial representatives in the family of crab spiders (Thomisidae) are a special case, as they build protective communal leaf nests instead of extensive communal capture webs. It could thus be inferred that antipredator benefits (e.g., enhanced protection in larger nests) rather than foraging‐related advantages (e.g., capture of more and larger prey) promote sociality in this family. Nonetheless, subsocial crab spiders do share prey, and if this behaviour does not reflect mere food scramble but has a cooperative character, crab spiders may offer insights into the evolution of social foraging applicable to many other cooperative predators that hunt without traps. Here, we performed a comparative laboratory feeding experiment on three of the four subsocial crab spider species—Australomisidia ergandros, Australomisidia socialis and Xysticus bimaculatus—to determine if crab spiders derive advantages from foraging in groups. In particular, we tested artificially composed groups of five sibling spiderlings vs. single siblings in terms of prey capture success and prey size preference. Across species, groups had higher prey capture success (measured in terms of capture rates and capture latency) and were more likely to attack large, sharable prey—dynamics leading to reduced food competition among group members in favour of living and foraging in groups. Within groups, we further compared prey extraction efficiency among the three applied social foraging tactics: producing, scrounging and feeding alone. In A. ergandros, individuals were exceptionally efficient when using the non‐cooperative scrounger tactic, which entails feeding on the prey provided by others. Thus, our multispecies comparison confirms foraging advantages in maintaining a cooperative lifestyle for crab spiders, but also demonstrates the relevance of research into exploitation of cooperative foraging in this family.     

Skills,division of labour and economies of scale among Amazonian hunters and South Indian honey collectors

In foraging and other productive activities, individuals make choices regarding whether and with whom to cooperate, and in what capacities. The size and composition of cooperative groups can be understood as a self-organized outcome of these choices, which are made under local ecological and social constraints. This article describes a theoretical framework for explaining the size and composition of foraging groups based on three principles: (i) the sexual division of labour; (ii) the intergenerational division of labour; and (iii) economies of scale in production. We test predictions from the theory with data from two field contexts: Tsimane'' game hunters of lowland Bolivia, and Jenu Kuruba honey collectors of South India. In each case, we estimate the impacts of group size and individual group members'' effort on group success. We characterize differences in the skill requirements of different foraging activities and show that individuals participate more frequently in activities in which they are more efficient. We evaluate returns to scale across different resource types and observe higher returns at larger group sizes in foraging activities (such as hunting large game) that benefit from coordinated and complementary roles. These results inform us that the foraging group size and composition are guided by the motivated choice of individuals on the basis of relative efficiency, benefits of cooperation, opportunity costs and other social considerations.     

Dominance and feeding interference in small groups of blackbirds

Dominance and/or interference parameters play a pivotal rolein most ideal free distribution models, but there has beenscant empirical study of the exact manner in which they jointlyoperate. We investigate how foraging effort and success variedamongst individuals of different dominance rankings in groupsof 1-3 wild blackbirds (Turdus merula) attracted to patchesof hidden food. Foraging effort (number of feeding movementsper unit time), as opposed to vigilance tradeoffs, was greaterwhen an individual fed with a subordinate conspecific thanwhen it fed alone, but tended to be less when it fed with adominant individual. Within dyads, changes in foraging effortwere associated with the direction of the dominance relationship,but not the relative difference in dominance rank between thetwo individuals. Similarly, amongst threesomes, top-rankedbirds (but not the lowest-ranked individual) showed higherforaging effort compared to when foraging alone. Top-ranked birds also profited from a greater increase in foraging success(food items per unit effort) than bottom-ranked birds whenfeeding in threesomes than when feeding alone. Dominant birdsshowed increased foraging success, but not effort, after displacinga subordinate. Our results suggest that an individual's foragingeffort is determined by the interplay of group vigilance benefitsand interference costs, the latter being more expensive for subordinate individuals. The foraging success of dominant birdsmay further increase if they use subordinates as food-finders.We discuss the implications of our findings for interferenceparameters in current Ideal Free Distribution models.     

Social information, social feeding, and competition in group-living goats (Capra hircus)

There are both benefits (e.g., social information) and costs(e.g., intraspecific competition) for individuals foraging ingroups. To ascertain how group-foraging goats (Capra hircus)deal with these trade-offs, we asked 1) do goats use socialinformation to make foraging decisions and 2) how do they adjusttheir intake rate in light of having attracted by other groupmembers? To establish whether goats use social information,we recorded their initial choice of different quality food patcheswhen they were ignorant of patch quality and when they couldobserve others foraging. After determining that goats use socialinformation, we recorded intake rates while they fed alone andin the presence of potential competitors. Intake rate increasedas the number of competitors increased. Interestingly, lonegoats achieved an intake rate that was higher than when onecompetitor was present but similar to when two or more competitorswere present. Faster intake rates may allow herbivores to ingesta larger portion of the available food before competing groupmembers arrive at the patch. This however, does not explainthe high intake rates achieved when the goats were alone. Weprovide 2 potential explanations: 1) faster intake rates area response to greater risk incurred by lone individuals, theloss of social information, and the fear of being left behindby the group and 2) when foraging alone, intake rate is no longera trade-off between reducing competition and acquiring socialinformation. Thus, individuals are able to feed close to theirmaximum rate.     

Risk Sensitivity and the Paradox of Colonial Web-Building in Spiders

Group foraging is rare in spiders, occurring only where preyavailability is high. If colonial web-building increases individualprey capture rates as shown, why does group foraging not occurmore often where prey are scarce? Risk sensitivity may explainthis paradox, as variance in prey capture is reduced in groups;risk-averse spiders should join groups only when prey exceeda threshold level. Field studies show that group foraging variesas predicted between species, between populations of a singlespecies, and between sites within a population. However, recentmodels suggest the necessity of examining variance within individualsover time rather than between individuals within populations.Additionally, mechanisms responsible for variance reductionin colonial webs may be less effective than previously assumed.New field data suggest that while prey variance over time maybe somewhat less for individual spiders in groups than for solitaries,the relationship between colonial web-building and variancein prey capture is far more complex than originally thought.The influence of risk sensitivity on reproductive success andthe evolution of colonial web-building is discussed.     

Number of neighbors instead of group size significantly affects individual vigilance levels in large animal aggregations

The group size effect states that animals living in groups gain anti‐predator benefits through reducing vigilance levels as group size increases. A basic assumption of group size effect is that all individuals are equally important for a focal individual, who may adjust its vigilance levels according to social information acquired from them. However, some studies have indicated that neighbors pose greater influences on an individual's vigilance decisions than other group members, especially in large aggregations. Vigilance has also been found to be directed to both predators (anti‐predation vigilance) and conspecifics (social vigilance). Central individuals might rely more on social vigilance than peripheral individuals. To test these hypotheses, we examined the effects of flock size, number of neighbors and position within a flock on vigilance and competition of greater white‐fronted goose Anser albifrons that form large foraging flocks in winter, controlling the effects of other variables (group identity, winter period and site). We found that individual vigilance levels were significantly affected by number of neighbors and position within a flock, whereas flock size showed no effect. Individuals devoted a large component of vigilance to nearby flock mates. Central individuals directed a relatively larger proportion of vigilance to monitor neighbors than peripheral ones, indicating that central individuals more relied on social information acquired from neighbors, possibly caused by the more blocked visual field of central individuals. Moreover, some social vigilance may function as conducting or preventing agonistic interactions since competition intensity was positively correlated with number of neighbors. Our study therefore demonstrate that the number of neighbors is more important than group size in determining individual vigilance in large animal groups. Further studies are still needed to unravel which neighbors pose greater influence on individual vigilance, and the factors that influence individuals to acquire information from their neighbors to adjust vigilance behaviors.     

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.     

Vigilance and Collective Detection of Predators in Degus (Octodon degus)

Individuals of social and partially social species typically reduce their vigilance activity when foraging in groups. As a result, per capita risk of predation decreases and individuals allocate more time to foraging and other fitness rewarding activities. Reduction of per capita risk is hypothesized to occur because there are more individuals to detect potential predators. If so, collective (i.e. total) vigilance is expected to increase with foraging group size. Increased surveillance during group foraging may occur if group members scan independently of one another, or sequentially to avoid the overlapping of their vigilance bouts. Intriguingly, such coordinated vigilance assumes that individuals monitor not only the presence, but the vigilance behaviour of group mates. We used seasonal records on time budget and grouping patterns of individually marked degus (Octodon degus), a social rodent, to examine if (a) individual vigilance decreases and/or foraging increases with group size, (b) collective vigilance increases with group size and (c) foraging degus coordinate their vigilance. When foraging, degus decreased their individual vigilance and increased their foraging time when in larger groups. Despite this, degus in larger groups increased their collective vigilance, supporting the hypothesis that socially foraging degus decrease predation risk through an improved ability to detect and escape potential predators. Additionally, patterns of collective vigilance suggested that degus scan independently of each other and so, they do not coordinate their vigilance to prevent its temporal overlapping. This finding does not support that foraging degus monitor the vigilance activity of group mates.     

Personality composition is more important than group size in determining collective foraging behaviour in the wild

Describing the factors that shape collective behaviour is central to our understanding of animal societies. Countless studies have demonstrated an effect of group size in the emergence of collective behaviours, but comparatively few have accounted for the composition/diversity of behavioural phenotypes, which is often conflated with group size. Here, we simultaneously examine the effect of personality composition and group size on nest architecture and collective foraging aggressiveness in the social spider Stegodyphus dumicola. We created colonies of two different sizes (10 or 30 individuals) and four compositions of boldness (all bold, all shy, mixed bold and shy, or average individuals) in the field and then measured their collective behaviour. Larger colonies produced bigger capture webs, while colonies containing a higher proportion of bold individuals responded to and attacked prey more rapidly. The number of attackers during collective foraging was determined jointly by composition and size, although composition had an effect size more than twice that of colony size: our results suggest that colonies of just 10 bold spiders would attack prey with as many attackers as colonies of 110 ‘average’ spiders. Thus, personality composition is a more potent (albeit more cryptic) determinant of collective foraging in these societies.     

东方田鼠社群气味对家族群成员个体觅食行为的影响

东方田鼠家族群成员个体的觅食行为是否因食物斑块存有家族群自身及非亲缘家族群气味而发生变异,进而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,分别配置家族群自身巢垫物及非亲缘家族群巢垫物作为社群气味,测定东方田鼠家族群在食物斑块觅食时,其成员个体觅食行为的序列过程及参数,检验家族群自身气味及非亲缘家族群气味对成员个体觅食行为的影响。结果表明,家族群自身气味能显著地缩短本群成员个体的觅食决定时间,通过减少成员个体的嗅闻及直立扫视动作时间比例、增大一般扫视、盯视及静听动作时间比例,降低觅食中断时间比例,提高其摄入率;而非亲缘家族群气味则能显著地延长家族群成员个体的觅食决定时间,通过增大家族群成员个体的嗅闻和一般扫视动作时间比例、减小直立扫视、盯视及静听动作时间比例,增大觅食中断时间比例,降低其摄入率。结果揭示,熟悉的社群气味会促使觅食活动中的家族群成员个体,在监测环境风险时,将精力更多地用于观察和监听群内其他成员个体的行为及其发出的警报信息,以便在有效规避环境风险的同时减缓个体间因干扰性竞争对觅食活动所造成的不利影响;而陌生的社群气味会迫使成员个体,将精力由依赖群内其他成员个体的行为...     

Social foragers adopt a riskier foraging mode in the centre of their groups

Foraging in groups provides many benefits that are not necessarily experienced the same way by all individuals. I explore the possibility that foraging mode, the way individuals exploit resources, varies as a function of spatial position in the group, reflecting commonly occurring spatial differences in predation risk. I show that semipalmated sandpipers (Calidris pusilla), a social foraging avian species, tended to adopt a riskier foraging mode in the central, more protected areas of their groups. Central birds effectively used the more peripheral group members as sentinels, allowing them to exploit a wider range of resources within the same group at the same time. This finding provides a novel benefit of living in groups, which may have a broad relevance given that social foraging species often exploit a large array of resources.     

Express your personality or go along with the group: what determines the behaviour of shoaling perch?

Behavioural syndromes, defined as correlated behaviours in different contexts, have been studied across species and taxa including humans as part of a personality concept. While most studies have focused on solitary individuals, less is known on how shoaling fish compromise between own personality and group behaviour. Risk-taking behaviour in 1-year-old perch (Perca fluviatilis) was observed to compare individual behaviour when in a group and when alone. An experimental design gave the fish the choice between foraging in an open area in the presence of a piscivore and hiding in the vegetation. We quantified the variation accountable by the effect of individuals being in a group, individuals alone and repeated measurements, using hierarchical mixed effects models. Within-group variances were low, but when individuals were later tested alone, individual differences explained most of the variation. Still, the individual best linear unbiased predictors (BLUPs) of time spent in the open area, extracted from the random effects of the mixed effects model, were positively correlated with the corresponding BLUPs when alone. The results indicate that individual behavioural traits are to some degree expressed also within groups. Most fish showed a shyer behaviour when alone, but bolder individuals changed less between treatments than did shyer ones, suggesting a more influential role of bold fish in the group.     

Evolution of group size in the dolphins and porpoises: interspecific consistency of intraspecific patterns

I investigated group size variability in dolphins and porpoisesusing intraspecific comparisons. Explanatory factors consideredin the analysis were variables of the physical environment,the diet, and the life history of the species. Open habitatand small body size were viewed as increasing predation risk.This pattern was apparent in Risso's dolphins (Grampus griseus) and weakly apparent in bottlenose dolphins (Tursiops spp.).Group size was negatively correlated with body size in pilotwhales (Globicephala spp.) and positively correlated with theopenness of habitat in killer whales (Orcinus orca), stripeddolphins (Stenella coeruleoalba), and common dolphins (Delphinus spp.). No such relationship was found for harbor and Dall'sporpoises (Phocoena phocoena, P. dalli). Group size also seemedto vary depending on other physical measures of the habitat,which may indirectly reflect diet; group size showed U-shapedpatterns if related to temperature. The predictive power ofvariables comprising detailed prey information on group sizewas variable. For example, pilot whales had smaller group sizes when they fed more on mesopelagic fish and less on mesopelagiccephalopods, and common dolphins had larger groups if theyfed on varying types of fish. In most Delphinoidea species,group size could be described by the variables considered inthis study. But each species showed its own pattern of correlationsbetween group size and a specific set of explanatory variables. Thus, no general and consistent relation between group sizeand the other variables was found. It remains unknown whetherthese species-specific patterns result from a historical processor whether they are specialized adaptations.     

Area-restricted search by the plains pocket gopher (Geomys bursarius) in tallgrass prairie habitat

Because pocket gophers have the high energetic cost of excavatingburrows and an inability to detect distant food items throughthe soil, I hypothesized that individuals within establishedburrow systems would use area-restricted search as a foragingstrategy. To examine this hypothesis I compared gopher foragingeffort over a 10-month period between areas in which overallplant densities were experimentally varied. Gophers expendedapproximately 50% of their foraging effort in areas with thehighest plant density, even though these made up only 33% ofthe available area in experimental plots. In large, griddedareas sampled for an entire season as well as in small areasin which gophers foraged for less than 1 week, gopher foragingeffort was related to the density of a single leguminous plantspecies, Psoralea argophylla. In small plots where this plantspecies was at high density, gophers created more tunnel branches,thereby intensifying their search effort. Thus, area-restrictedsearch appears to increase the rate of encounter with the patchilydistributed Psoralea plants.     

The benefits of stealing from a predator: foraging rates, predation risk, and intraspecific aggression in the kleptoparasitic spider Argyrodes antipodiana

In the trade-off between food and safety, the role of aggressiveintraspecific interactions has not been extensively examined.Here I present information on this system using a kleptoparasiticspider, Argyrodts antipodiana, and its host spider and potentialpredator, Eriophora pustulosa. A. antipodiana can feed eitherat a potentially dangerous site (the hub of its host's web withthe host), or at a relatively safe site (on food bundles aroundthe edge of the host's web). I found that A. antipodiana cangain food very quickly when feeding with the host, apparentlyby exploiting the host's ability to digest the prey. Thus A.antipodiana follows predictions based on foraging models inthat it accepts a higher predation risk at the hub because ofthe higher food payoff. A. antipodiana also aggressively competesfor access to more food. However, aggressive competition increasesthe predation risk from the host, especially at tile hub wherethe host is very close. Consequently, A. antipodiana modifiesits level of intraspecific aggressiveness in accordance withits position on the web: at the hub, where the cost of aggressionis high (due to predation risk), A. antipodiana reduces itsaggressiveness, but it is aggressive away from the hub whencompeting for food bundles. The ability of A. antipodiana tochange interaction intensity as a function of its position onthe web enables it to exploit a rich, but risky, food sourceand provides a new angle for examining food and safety tradeoffs in light of intraspecific competition for food     

Resource patch size and flexible foraging in white-faced capuchins (Cebus capucinus)

White-faced capuchins (Cebus capucinus)on Barro Colorado Island, Panama, have a flexible foraging strategy. Typically, foraging party size is small and individuals feed dispersed from one another. When seasonal fruiting of large volume trees occurs, the majority of the group forages simultaneously. As C. capucinusdo not display a rigorous dominance structure and there are few indications that individuals or coalitions monopolize food patches,individuals are expected to display scramble strategies instead of high frequencies of contest competition. I recorded foraging party size (simultaneous foragers), the total number of animals to feed successively, and the diameter at breast height (DBH) of fruit trees used in two habituated troops. Individuals in each group spent a substantial amount of time — 65 and 48% of foraging time for each group — foraging in party sizes of one. Monkeys predominantly foraged alone in small trees (0- to 20- cm DBH), successively in medium trees (21- to 60- cm DBH), and simultaneously in large trees (>61- cm DBH). They used small trees more frequently than all other tree classes. In medium-sized trees, although fruit was plentiful, space was limited. In these trees Cebusforaged successively. In large-volume trees, space and fruit were abundant and several individuals fed together. As the DBH of fruiting trees increased, the average foraging party size increased exponentially. Cebus capucinusat Barro Colorado Island modify their foraging party size to adapt to the seasonal patterns of fruit production.