Do kleptoparasites reduce their own foraging effort in order to detect kleptoparasitic opportunities?: An empirical test of a key assumption of kleptoparasitic models

Determining if, or when, individuals trade off time spent personally feeding against time spent monitoring others for kleptoparasitism opportunities is essential to an understanding of the evolution of scrounging and usurpation behaviours. We provide a first field test of whether kleptoparasites reduce their personal foraging effort in situations where the frequency and rewards of kleptoparasitism increase. We provided experimental food patches for wild European blackbirds that varied in the distribution of prey and that had a potentially high rate of kleptoparasitism within pairs of blackbirds feeding in them. Although individuals differed in their rate of kleptoparasitism, they did not vary in the size of the reward that they gained from kleptoparasitism. As prey became more clumped, kleptoparasitism rate and its reward per incident increased on average. There was, however, no evidence that individuals that were kleptoparasitising more quickly and/or at a higher frequency had lower personal foraging effort. In contrast, foraging effort increased in both birds compared to when they were foraging alone, independent of dominance, kleptoparasitic opportunity or reward. Our evidence suggests that in some circumstances a kleptoparasite can detect kleptoparasitic opportunities without compromising its own personal foraging rate.     

Movement coordination and signalling in ravens (Corvus corax): an experimental field study

Vagrant non-breeding ravens frequently attract conspecifics to rich ephemeral food sources. There, grouping may allow them to overcome the defence of territorial breeders. Here, we focus on ravens making use of regular food supplies in a game park, where they divert food from the provision of park animals. We investigated if ravens foraging in the Cumberland game park (Grünau, Austria) are attentive towards one another when they experience some unpredictability in food provisioning. We confronted a group of 30–50 ravens with two different treatments. Ten minutes ahead of the feeding of either wolves or wild boars we showed buckets containing pieces of meat to the ravens flying overhead. In the reliable cue treatment (RCT), the meat was placed next to one of the two enclosures, whereas in the unreliable cue treatment (UCT), the buckets were placed simultaneously in front of both enclosures though only in one of the enclosures were the animals fed 10 min later. Thus, during RCT but not during UCT, ravens could predict where food would become available. Only during UCT, ravens moved in large groups between the two feeding sites. Many ravens moving at the same time in the same direction may indicate some co-ordination in space and time, which is most likely achieved by social attraction among individuals. Furthermore, the number of ravens approaching and leaving, respectively, a feeding site cross-correlated with a temporary increase in the rate of a food-associated call, the yell. This suggests that in addition to watching each other, calling may have contributed to group formation. Possible benefits of group formation during food inspection are discussed. Received: 9 February 2000 / Received in revised form: 30 May 2000 / Accepted: 1 June 2000     

Common ravens, Corvus corax, preferentially associate with grey wolves, Canis lupus, as a foraging strategy in winter

One foraging strategy that scavengers can employ to discover unpredictable food sources is to associate directly with predators who inadvertently provide food. The common raven, a well known feeding generalist, is also a prominent scavenger of wolves' kills and is found to be in close association with this predator. We tested the hypothesis that ravens preferentially associate with wolves in winter as a kleptoparasitic foraging strategy. The presence, absence and behaviour of ravens was documented during winter observations of wolves, coyotes, Canis latrans, and elk, Cervus elaphus, as well as the landscape in the absence of these three species. Ravens were found to be in close association with wolves when they were travelling, resting and hunting prey. In comparison, ravens showed no significant association with coyotes, elk or areas on the landscape in the absence of wolves. We also compared ravens' discovery success of wolf-killed and nonwolf-killed carcasses and their behavioural response upon discovery. Ravens found all wolf kills almost immediately and remained at the carcass to feed alongside wolves after the death of the prey. In contrast, ravens were less successful discovering experimentally placed carcasses in the same study region, and did not land or feed despite the availability of fresh, exposed meat. Our results show that ravens' association with wolves is not just an incidental and proximate by-product of the presence of fresh meat. Instead, we show that ravens preferentially associate with wolves in both the presence and absence of food, resulting in the discovery of carcasses and suppression of ravens' innate fear of novel food sources. Through this mode of social foraging, ravens may experience increased foraging efficiency in the use of an otherwise spatially and temporally unpredictable food source.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Effect of Group Size on Feeding Rate when Patches are Exhaustible

One benefit of group foraging is that individual foragers can join the food discoveries of companions and thus increase encounter rate with food patches. When food patches are exhaustible, however, individual shares of each patch will decrease with group size negating the effect of increased encounter rate. Mean feeding rate may actually decrease with group size as a result of aggression or time wasted joining already depleted patches, or when searching to join the food discoveries of others, which is referred to as scrounging, precludes finding food. I examined the relationship between mean feeding rate and group size in captive flocks of zebra finches (Taenopygia guttata) foraging for small clumps of seeds. Finches in groups of two or four fared better than solitary birds in terms of mean feeding rate despite the fact that birds in groups scrounged a large proportion of their food. Solitary birds initiated feeding activity after a longer delay, which led to their lower success. Early departures by food finders from food patches joined by others may have lessened the impact of scrounging on mean feeding rate. As a result of benefits from the presence of companions, group foraging in zebra finches appears a viable alternative to foraging alone despite the cost of sharing resources.     

The Effect of Within-Flock Spatial Position on the Use of Social Foraging Tactics in Free-Living Tree Sparrows

The benefit of producer (searches for own food) or scrounger (exploits the others’ food discoveries) foraging tactic in a group of socially feeding animals may depend on where the individual searches for food within the group. Scrounging may be more advantageous in the centre of the group, having more individuals around to join, while producing may be more beneficial at the edges, where more unexplored food patches may be found. This study shows within‐flock position correlates with foraging tactic use of feeding birds in socially foraging tree sparrows, Passer montanus. Sparrows staying closer to the centre of the flock found their food patches more frequently by joining (i.e. use more frequently the scrounging tactic) than those staying toward the edges. To our knowledge this is the first field study demonstrating the relationship between spatial position and foraging tactic use. We investigated this relationship under different perceived predation hazard, and found that under elevated risk of predation, central individuals may increase their use of joining more than individuals on the periphery of the flock. Moreover, we show that extremely specialized use of searching tactics may be very infrequent in tree sparrows. As both within‐flock position and search tactic use can be altered very quickly and without leaving the flock, individuals may easily alter them in order to adjust their behaviour.     

Phenotypic Correlates of Scrounging Behavior in Zebra Finches: Role of Foraging Efficiency and Dominance

In flocks, individuals can search for their own food using the producer tactic or exploit the discoveries of companions using the scrounger tactic. Models of the producer–scrounger game usually assume that tactic payoffs are independent of individual phenotypic traits. However, factors such as dominance status or foraging efficiency may constrain the use of tactics and lead to asymmetric tactic use among individuals. For instance, in flocks composed of foragers with unequal foraging efficiency, foragers that are less efficient at obtaining food are expected to rely on the scrounger tactic to a greater extent. I examined the role of foraging efficiency and dominance status as potential correlates of scrounging behavior in small aviary flocks of zebra finches (Taenopygia guttata). Individual foraging efficiency was documented in each flock in a treatment that prevented scrounging. In a subsequent treatment that allowed scrounging, higher levels of scrounging occurred as predicted in foragers with lower foraging efficiency. Dominance status was a poor predictor of tactic choice. Birds that arrived later on the foraging grid foraged less efficiently when scrounging was prevented and used scrounging to a greater extent when allowed, suggesting a link between boldness, foraging efficiency and the choice of foraging tactics in small flocks of zebra finches.     

Ravens adjust their antipredatory responses to con‐ and hetero‐specific alarms to the perceived threat

Heterospecific alarm calls are typically found in situations where multiple species have a common predator. In birds, they are particularly common in mixed mixed‐species flocks. In species with highly developed social and cognitive abilities like corvids, there is the potential for differential responses to heterospecific vs. conspecific calls according to the riskiness of the habitat. We tested the responses of free‐ranging ravens (Corvus corax) to conspecific alarm calls and compared them to heterospecific alarm calls of jackdaws (Corvus monedula). We observed the proportion of ravens leaving the feeding site after the con‐ or hetero‐specific playback was presented in a situation of low threat (wild boar—Sus scrofa enclosure) and high threat of predation (wolf—Canis lupus enclosure). We show that ravens responded to conspecific calls more intensively at the wolves than at the wild boar, but the response to conspecific calls was in both enclosures stronger than to the control (great tit—Parus major song). The response to the heterospecific alarm was also stronger in the wolves’ enclosure, but it did not differ from control in the wild boar enclosure. These findings suggest that ravens are aware of the meaning of the jackdaw alarm calls, but they respond to it only in a situation of high predatory threat (wolves are present). In the wild boar enclosure, the ravens probably consider jackdaws warning against some other predator, very probably harmless to ravens. This interpretation requires further testing, as both enclosures differ also in respect to other parameters like food quality and shelter availability.     

Neophobia affects choice of food-item size in group-foraging common ravens (Corvus corax)

Individuals foraging in groups should develop behavioural tactics to optimise their gain. In novel feeding situations, predation risk and pressure of kleptoparasites may be particularly high and hence may constrain optimal foraging. To create a novel feeding situation, we offered common ravens (Corvus corax) equal numbers of either small (40 g) or large (160 g) pieces of meat on successive days, always in combination with the same novel object. During the first weeks, when ravens were still neophobic, small pieces were taken in larger numbers than large pieces. Intraspecific kleptoparasitism was more likely to occur when ravens carried large food items. It seems that initiating foragers were mainly innovative subdominants. Preference for small items might have decreased with increasing habituation because more dominants were then feeding directly at the source and hence were less likely to resort to kleptoparasitism as an alternative foraging tactic. Electronic Publication     

Wild Carib grackles play a producer scrounger game

Producer–scrounger (PS) game-theoretical foraging modelsmake predictions about the decision of group-feeding animalseither to look for food (produce) or for opportunities to exploitthe discoveries of other foragers (scrounge). We report themost complete demonstration to date of the applicability ofthe PS foraging game in a free-living animal, the Carib grackle(Quiscalus lugubris) of Barbados. As assumed by PS games, thepayoffs obtained by scroungers were negatively frequency dependent.Experimentally, increasing the cost of scrounging led to a decreasein the observed proportion of scroungers, whereas raising thecost of producing increased the proportion of scroungers. Observationsof marked birds revealed that group-level changes could be broughtabout by individual flexibility in tactic use. Despite consistentindividual differences in tactic use, most birds used both tacticsand could alter their use of producing and scrounging when conditionschanged. We found no difference in the payoffs obtained by producersand scroungers, suggesting a symmetrical game equilibrium. Ourresults call for testing the PS foraging game in a broader rangeof biological systems that include different types of scroungingbehavior (e.g., scramble, stealthful, or aggressive scrounging)as well as the exploitation of different phases of food production(e.g., searching, handling).     

The effects of water on patch use by two Simpson Desert granivores (Corvus coronoides and Pseudomys hermannsburgensis)

Abstract Water loss while foraging may affect the overall value of food to desert animals. When water is scarce, foragers may alter activity and shun certain types of food due to elevated water loss. When water is abundant, foragers can exploit food patches more thoroughly and remain active over a broader range of ambient conditions. In short, food and water may be complementary resources. The presence of water raises the marginal value of food, particularly those foods low in water content. We tested for the complementarity of food and water to foragers at a sand dune site in the Simpson Desert of arid Australia. To do so, we quantified patch exploitation of foragers in the presence or absence of bowls filled with water. In order to quantify patch use, we provisioned feeding trays with granulated peanuts mixed into a sand substrate. In these trays we measured giving-up densities (GUD; the amount of food left in a tray after a foraging bout) of diurnal (mostly Australian ravens, Corvus coronoides) and nocturnal foragers (mostly sandy inland mouse, Pseudomys hermannsburgensis). The presence of water affected the GUD of ravens but not of rodents. For the ravens, GUD dropped about 50% in response to added water. For ravens, water and food are strongly complementary. In addition, ravens had lower GUD in the open than the bush microhabitat, and lower GUD at the bottom than the tops of sand dunes.     

Raven scavenging favours group foraging in wolves

Wolves, Canis lupus, routinely live in large packs that include unrelated individuals and mature offspring. Studies show that individual wolves that live in large packs suffer reduced foraging returns. Therefore, group hunting and group living (sociality) in wolves is generally thought to be favoured by indirect fitness gains accrued through kin-directed altruism. However, we show that kin-directed altruism cannot account for groups that include mature offspring or unrelated individuals. We also present an analysis that incorporates a previously ignored feature of wolf foraging ecology, namely the loss of food to scavenging ravens, Corvus corax. By accounting for this process, we show that individuals in large packs do indeed accrue foraging advantages. In the hypothetical absence of this scavenging pressure, an individual would maximize its rate of prey acquisition, and minimize its risk of energetic shortfall, by foraging with just one other individual. However, incorporating the effect of scavenging by ravens leads to a dramatic increase in the predicted group size. Our analysis indicates that per capita gains are highest in the largest observed packs. The greater food-sharing costs in a larger pack are more than offset by smaller losses to scavengers and increased rates of prey acquisition. Thus, in contrast with previous interpretations, the selfish benefits of social foraging appear to contribute to the maintenance of sociality in wolves after all. We explore whether such benefits favour group living in various social carnivores that hunt large prey and are thus vulnerable to scavenging.     

Decision time modulates social foraging success in wild common ravens,Corvus corax

Social foraging provides several benefits for individuals but also bears the potential costs of higher competition. In some species, such competition arises through kleptoparasitism, that is when an animal takes food which was caught or collected by a member of its social group. Except in the context of caching, few studies have investigated how individuals avoid kleptoparasitism, which could be based on physical strength/dominance but also cognitive skills. Here, we investigated the foraging success of wild common ravens, Corvus corax, experiencing high levels of kleptoparasitism from conspecifics when snatching food from the daily feedings of captive wild boars in a game park in the Austrian Alps. Success in keeping the food depended mainly on the individuals’ age class and was positively correlated with the time to make a decision in whether to fly off with food or consume it on site. While the effect of age class suggests that dominant and/or experienced individuals are better in avoiding kleptoparasitism, the effect of decision time indicates that individuals benefit from applying cognition to such decision-making, independently of age class. We discuss our findings in the context of the ecological and social intelligence hypotheses referring to the development of cognitive abilities. We conclude that investigating which factors underline kleptoparasitism avoidance is a promising scenario to test specific predictions derived from these hypotheses.     

Measuring social tolerance: An experimental approach in two lemurid primates

Social tolerance crucially affects the life of group‐living animals as it can influence, among other things, their competitive regimes, access to food, learning behavior, and recruitment. However, social tolerance tests were mainly conducted in semi‐free or captive populations, and we know little about the behavioral mechanisms and consequences of social tolerance under natural conditions. We therefore developed a co‐feeding experiment to measure social tolerance in groups of wild and captive animals across two primate species. Specifically, we recorded the social tolerance level of redfronted lemurs (Eulemur rufifrons, four wild, one captive group) and ringtailed lemurs (Lemur catta, three wild, three captive groups) by presenting a clumped food resource in an experimental arena, and recorded patterns of resource use during the experiment. Because redfronted lemurs exhibit lower levels of decided conflicts than ringtailed lemurs, we predicted that they would be socially more tolerant. The probability for an individual to feed in the arena was higher in redfronted lemurs than in ringtailed lemurs. In addition, in both species, the probability for an individual to feed in the arena was higher in the captive populations than in their wild counterparts, suggesting that proximate factors, such as a relaxation of feeding competition in captivity, may adapt species‐specific levels of social tolerance to local levels of food availability. Hence, the number of individuals co‐feeding on a valuable food resource appears to be a useful proxy of social tolerance that could be measured with this experimental setup in other wild and captive species as well.     

Food calling in wild ravens (Corvus corax) revisited: Who is addressed?

Numerous birds and mammals use vocal signals to advertise feeding opportunities but often such signals vary with individual and contextual factors. Non-breeding ravens call at food that is difficult to access, resulting in the attraction of nearby conspecifics. Although callers may benefit from group formation in various ways, we recently found substantial individual variation in food calling. We here explored whether this variation can be partly explained by the social dynamics in raven foraging groups, together with already known effects of age class and sex. Specifically, we expected ravens to respond to the presence or absence of affiliates that could act as cooperative partners in the forthcoming feeding event, that is they should call when other ravens were present but they themselves were alone rather than when they were also in company of an affiliation partner. We observed the vocal behaviour of individually marked wild ravens and, simultaneously, categorized their affiliative behaviour with other ravens in the minutes before experimentally controlled feedings. In line with our prediction, individuals were less likely to produce food-associated calls when they were in close contact with an affiliation partner prior to feeding as compared to when they were alone. Furthermore, sex and age class influenced food calling as females called more often than males and younger birds called more often than adult ravens. In conclusion, these results suggest that ravens attempt to find support from a particular cooperative partner by broadly advertise feeding opportunities via food-associated calls, especially when they have low chances in contest competition due to their age and sex. These findings lend further support to the assumption of raven flocks being structured by social relationships and individual birds flexibly controlling their vocal signalling according to the current flock composition.     

Habitat selection at multiple spatial scales by foraging Glossy Black‐cockatoos

Abstract: Habitat selection among vertebrates entails decision making at a number of spatial scales. An understanding of factors influencing decisions at each of these scales is required for the effective management of wildlife populations. This study investigates the foraging ecology of a population of Glossy Black‐cockatoos in central New South Wales. We took advantage of the characteristic feeding sign produced by Glossy Black‐cockatoos to examine factors influencing habitat selection at multiple spatial scales. Birds preferred to forage at sites where food was abundant and avoided open sites where the predation risk may be greater. Their two food species, Allocasuarina diminuta and Allocasuarina gymnanthera, differed in profitability (kernel intake rate as measured by the ratio of seed weight to total seed and cone weight), as did trees within a species. Both species were utilized extensively, although foraging intensity was greater at sites where the more profitable species was present. In order to maximize their food intake, birds selected individual trees on the basis of cone abundance and profitability. Cones produced in the previous year were preferred.     

Individual variation in the foraging strategies of young yellow perch (Perca flavescens) from the Ottawa River

Differences in the foraging strategies among young individuals of the yellow perch (Perca flavescens) were observed in the laboratory by using two kinds of food (Daphnia and brine shrimp) separately and together. Individuals differed significantly in their ability for feeding attempts, time interval between two consecutive feeding attempts, feeding angles, regurgitation rate, and number of unsuccessful attempts and in their food preference. It is concluded that there are individuals with different foraging strategies. Variability in foraging strategies within single species populations is important because it may explain how the conspecific individuals may differ in their overall feeding behaviour.     

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.     

Experimental Manipulation of Food Accessibility Affects Conflict Management Behaviour in Ravens

Conflict management strategies such as reconciliation and bystander affiliation have been described for a variety of species. A common determinant seems to be a ‘complex’ social life, with individuals relying on affiliate relationships or social bonds. Little is known, however, about the strategic and flexible use of conflict management skills in experimental settings in species other than primates. We here investigated conflict and post‐conflict behaviour of ravens by manipulating the accessibility of food and, thus, the likelihood of aggressive interactions while foraging. Specifically, we presented birds with a certain amount of highly preferred food that varied in the number of pieces (one piece, two pieces or, as a control, small pieces matching the number of participating birds) and observed their agonistic behaviour during feeding and their affiliative behaviour afterwards. The results showed that high levels of conflicts during feeding in the 1‐piece and 2‐piece conditions led to high levels of affiliation after feeding. Depending on the experimental condition, this effect is best explained (a) by the affiliative behaviour of former aggressors (1‐piece condition) and (b) by the affiliation directed to the receivers of aggression after feeding (2‐piece condition). Those dyads that engaged in allo‐preening after feeding also engaged in allo‐preening outside the experimental setting, suggesting that socially bonded individuals provided third‐party affiliation to victims of aggression. Moreover, socially bonded ravens fed close to each other in the experiment when food was clumped, indicating that they actively coordinated their behaviour when there was a high conflict potential. Taken together, these findings support the assumption that ravens use their social bonds to avoid conflicts by choosing with whom to feed, and to buffer effects of conflicts by engaging in third‐party affiliation as post‐conflict behaviour.     

The feeding habits of wolves in relation to large prey availability in northern Italy

We investigated wolf feeding habits in relation to the abundance of wild and domestic ungulates to test the hypothesis that large prey are preferred and that their abundance affects the use of other food categories and diet breadth. We determined diet composition by scat analysis from December 1987 to December 1992. The research was carried out in three study areas located in northern Italy and characterised by marked differences in wild and domestic ungulate abundance. In study area A (low wild and domestic ungulate availability) fruits, livestock, other vertebrates and wild ungulates made up the bulk of the diet (71% in volume). In area B (high availability of livestock) wolf diet was mainly based on sheep and wild boars (80% in volume). In study area C (high availability of wild ungulates) wild ungulates were the main food of wolves (90% in volume). Significant differences were found among study areas in the mean percentage volume of all food categories and in particular for wild ungulates, livestock, other vertebrates and fruits (p < 0.0001 in all cases). Diet breadth decreased in areas with high availability of large wild and domestic herbivores. The use of livestock species was lower where there was high abundance, richness and diversity of the wild ungulate guild. Selection for wild ungulate species was partially affected by their abundance: however other factors as prey social behaviour, adaptability to the habitat (for introduced species), and body size could have an important role in species selection by wolves. In particular in area C wild boars were selected for, roe and red deers avoided, and fallow deers and mouflons used as available. Livestock species were used in relation to their abundance and accessibility, in particular sheep were selected for and cattle avoided; but if calves bom in the pastures were considered as the only available cattle, they were selected for and sheep were used as available. Large and in particular wild herbivores were found to be of great importance for the wolf population maintenance in northern Italy, one of the most important recovery areas of Mediterranean wolves.     

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.