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.     

Mixed species flocking of tits (Parus spp.): a field experiment

Summary We tested two general models of flocking behaviour, namely the antipredation model and foraging efficiency model on mixed-species tit flocks (Parus spp.). After food addition the size of mixed-species flocks was significantly less than in the control samples. In the presence of extra food significantly more birds were observed either in monospecific flocks or solitary, than during the control observations. In the presence of a living predator the birds foraged in larger mixed-specifies flocks than during the control observations. In addition, the social behaviour of Great Spotted Woodpecker, Middle Spotted Woodpecker and Nuthatch shifted to mixed-specific flocking. The size of monospecific flocks was independent of both treatments. The density of birds increased significantly after food addition, while in the predator presence the birds tended to leave the forest. These results support the view that both the antipredation model and foraging efficiency model seem to be valid for mixed-species flocking. However, in the case of monospecific flocks, the territory maintenance could be the most important factor.     

Bird mixed‐species flock formation is driven by low temperatures between and within seasons in a Subtropical Andean‐foothill forest

According to both the predation avoidance and foraging efficiency hypotheses, birds within mixed flocks increase their foraging efficiency and/or can spend more time feeding and less time looking out for predators. These hypotheses predict that birds in mixed flocks obtain benefits. Thus, mixed flock formation could serve as a strategy to cope with difficult conditions imposed on birds such as climatic conditions that ultimately result in a change in predation pressure or food resources. We evaluate the hypotheses that forming part of a flock confers benefits to its members and the associated prediction that birds will take advantage of these benefits and flock more often under cold and dry weather conditions between and within seasons to cope with such conditions. We surveyed the presence of mixed flocks, flocking propensity, number of species and individuals in mixed flocks in the Subtropical Yungas foothill of Argentina, to examine seasonality, flocking behavior of birds and their responses to two climatic variables: temperature and humidity. Bird species presented a higher flocking propensity and mixed flocks occurred more frequently during the dry and cold seasons than during the more benign seasons, and lower values of temperature within seasons triggered the flocking behavior. Although effects between seasons were expected, birds also showed a short‐term response to small changes in temperature within seasons. These results strengthen the ideas proposed by the foraging hypothesis. Although benefits derived from flocking have yet to be determined, whatever they are should be understood in the context of seasonal variation in life‐history traits.     

Effects of supplemental food on intra-and inter-specific behaviour of the Varied Tit Parus varius

Varied Tits Parus varius lived in flocks containing a pair, or a pair with one or two unpaired birds. To test whether the sociality of Varied Tits or their participation in mixed-species flocks is sensitive to additional food (sunflower seeds), we compared the following parameters between fed and unfed periods at the individual level: population size, degree of site fidelity, mono-specific flock size and the stability of its membership, home-range size and distribution, attendance rate with mixed-species flocks, and flock size. Neither the size of the mono-specific flock nor its membership was affected by food supply. Whether food was added or not, individuals exhibited a strong site fidelity resulting in stable population size. Supplemental feeding had no effect on home-range size or distribution. In the presence of extra food, Varied Tits were observed in mono-specific flocks but rarely in mixed-species flocks. However, when we stopped feeding, they shifted to mixed-species flocking. After adding food, mixed-species flocks were significantly smaller than in the control samples. Varied Tits were more likely to join mixed-species flocks as temperatures dropped and wind speed increased. These results suggest that intra-specific sociality of Varied Tits is relatively insensitive to food supply, but they easily shift to mixed-species flocking in relation to food and weather conditions. We conclude that Varied Tits participate in mixed-species flocks to obtain short-term benefits e.g. increased foraging efficiency but they also obtain long-term benefits from stability of pair bonds and strong site fidelity, which did not respond to supplemental food.     

Reduced flocking by birds on islands with relaxed predation

Adaptive hypotheses for the evolution of flocking in birds have usually focused on predation avoidance or foraging enhancement. It still remains unclear to what extent each factor has contributed to the evolution of flocking. If predation avoidance were the sole factor involved, flocking should not be prevalent when predation is relaxed. I examined flocking tendencies along with mean and maximum flock size in species living on islands where predation risk is either absent or negligible and then compared these results with matched counterparts on the mainland. The dataset consisted of 46 pairs of species from 22 different islands across the world. The tendency to flock was retained on islands in most species, but in pairs with dissimilar flocking tendencies, island species were less likely to flock. Mean and maximum flock size were smaller on islands than on the mainland. Potential confounding factors such as population density, nest predation, habitat type, food type and body mass failed to account for the results. The results suggest that predation is a significant factor in the evolution of flocking in birds. Nevertheless, predation and other factors, such as foraging enhancement, probably act together to maintain the trait in most species.     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

The effect of prior residence and pair bond on scrounging choices in flocks of zebra finches, Taenopygia guttata

In groups, animals can use the producer tactic to locate food patches and the scrounger tactic to join the food discoveries of other companions. At equilibrium, models predict a mixture of the two tactics with equal payoffs. Several factors may constrain the use of tactics and lead to biases in scrounging choices. I explored the effect of prior residence and pair bond as potential constraints on scrounging choices in flocks of zebra finches (Taenopygia guttata). Experimental flocks contained two birds already established in an aviary (prior residents) and two birds recently released in the aviary for the first time (new residents). All birds were previously trained to find food on a foraging grid. In the aviary, new residents followed prior residents from perches to the grid and relied heavily on prior residents to locate food patches. Low initial success by new residents probably favoured heavy reliance on the scrounger tactic. New residents that formed pair bonds with prior residents foraged closer to their mates and scrounged selectively from their mates in some cases. Prior residence, and pair bond to a lesser extent, influenced scrounging choices in zebra finches and could lead to deviation from the expected use of foraging tactics.     

Social networks predict patch discovery in a wild population of songbirds

Animals use social information in a wide variety of contexts. Its extensive use by individuals to locate food patches has been documented in a number of species, and various mechanisms of discovery have been identified. However, less is known about whether individuals differ in their access to, and use of, social information to find food. We measured the social network of a wild population of three sympatric tit species (family Paridae) and then recorded individual discovery of novel food patches. By using recently developed methods for network-based diffusion analysis, we show that order of arrival at new food patches was predicted by social associations. Models based only on group searching did not explain this relationship. Furthermore, network position was correlated with likelihood of patch discovery, with central individuals more likely to locate and use novel foraging patches than those with limited social connections. These results demonstrate the utility of social network analysis as a method to investigate social information use, and suggest that the greater probability of receiving social information about new foraging patches confers a benefit on more socially connected individuals.     

Tolerance of understory plants subject to herbivory by roe deer

Classical foraging theory states that animals feeding in a patchy environment can maximise their long term prey capture rates by quitting food patches when they have depleted prey to a certain threshold level. Theory suggests that social foragers may be better able to do this if all individuals in a group have access to the prey capture information of all other group members. This will allow all foragers to make a more accurate estimation of the patch quality over time and hence enable them to quit patches closer to the optimal prey threshold level. We develop a model to examine the foraging efficiency of three strategies that could be used by a cohesive foraging group to initiate quitting a patch, where foragers do not use such information, and compare these with a fourth strategy in which foragers use public information of all prey capture events made by the group. We carried out simulations in six different prey environments, in which we varied the mean number of prey per patch and the variance of prey number between patches. Groups sharing public information were able to consistently quit patches close to the optimal prey threshold level, and obtained constant prey capture rates, in groups of all sizes. In contrast all groups not sharing public information quit patches progressively earlier than the optimal prey threshold value, and experienced decreasing prey capture rates, as group size increased. This is more apparent as the variance in prey number between patches increases. Thus in a patchy environment, where uncertainty is high, although public information use does not increase the foraging efficiency of groups over that of a lone forager, it certainly offers benefits over groups which do not, and particularly where group size is large.     

Effects of Resource Allocation on Behavioural Strategies: A Comparison of Red Junglefowl (Gallus gallus) and Two Domesticated Breeds of Poultry

Three different breeds of poultry, representing different degrees of domestication, were observed in semi-natural conditions in order to study differences in foraging behaviour, activity levels and social behaviours which could be caused by correlated responses to selection for increased production. The breeds used were: (i) red junglefowl ( Gallus gallus ); (ii) Swedish bantam, which is a domestic breed that has not undergone selection for production traits; and (iii) Hy-Line, a White leghorn laying hybrid, selected mainly for food conversion efficiency. The birds were offered ad libitum food simultaneously from sites where the food was freely available and from sites where the birds had to search and scratch for food which was mixed with wood-shavings. The behaviour was observed three times per day (48 min/d), 3 d per week in eight groups (four birds per group) of each breed between 7 and 18 wks of age. Junglefowl and bantam obtained a significantly higher proportion of their food from the site that required effort. The opposite case was true for the Hy-Line. Overall, bantams performed significantly more foraging behaviour than Hy-Lines. The Hy-Line breed was more inactive and less involved in social interactions than the junglefowl and the bantam. The results support the idea that selection for high production results in modified behavioural strategies. Behaviours that are of high energetic cost, such as extensive foraging and social interactions, were of lower in frequency in the laying hens compared to junglefowl and bantam, allowing the laying hens to save energy that can be reallocated to production traits.     

Sex differences in giraffe foraging behavior at two spatial scales

We test predictions about differences in the foraging behaviors of male and female giraffes (Giraffa camelopardalis tippelskirchi Matchie) that derive from a hypothesis linking sexual size dimorphism to foraging behavior. This body-size hypothesis predicts that males will exhibit specific behaviors that increase their dry-matter intake rate relative to females. Foraging behavior was examined at two hierarchical levels corresponding to two spatial and temporal scales, within patches and within habitats. Patches are defined as individual trees or shrubs and habitats are defined as collections of patches within plant communities. Males were predicted to increase dry-matter intake rate within patches by taking larger bites, cropping bites more quickly, chewing less, and chewing faster. Within habitats, males were expected to increase intake rate by increasing the proportion of foraging time devoted to food ingestion as opposed to inter-patch travel time and vigilance. The predictions were tested in a free-ranging population of giraffes in Mikumi National Park, Tanzania. Males spent less total time foraging than females but allocated a greater proportion of their foraging time to forage ingestion as opposed to travel between patches. There was no sex difference in rumination time but males spent more time in activities other than foraging and rumination, such as walking. Within patches, males took larger bites than females, but females cropped bites more quickly and chewed faster. Males had longer per-bite handling times than females but had shorter handling times per gram of intake. Within habitats, males had longer average patch residence times but there was no significant sex difference in inter-patch travel times. There was no overall difference between sexes in vigilance while foraging, although there were significant sex by habitat and sex by season interactions. Although not all the predictions were confirmed, overall the results agree qualitatively with the body-size hypothesis. Sex-related differences in foraging behavior led to greater estimated intake rates for males at the within-patch and within-habitat scales. Received: 20 November 1995 / Accepted: 5 November 1996     

Communal Roosting of Wintering Red Kites Milvus milvus (Aves,Accipitridae): Social Feeding Strategies for the Exploitation of Food Resources

Social feeding strategies of wintering red kites are analyzed in relation to age, food, roost-sites and differences from kite residents. Whereas young and adult wintering kites gathered at roost sites almost daily, adult residents did not, and immature residents only occasionally. Kites using roost sites feed more often on prey prelocated by others, while lone roosters also forage and discover food alone. After finding food, kites tend to shift to a new roost site and foraging area. Two details of the ‘information centre’ hypothesis are confirmed in our study: carcasses are unpredictably found patches, divisible between several individuals. But carcasses disappeared fast in the study area, and no increase with time in the number of birds consuming a carcass was observed, so that information transmission was unconfirmed. When kites leave the roost in groups no leader is detectable. It seems that other types of social foraging are operating, and the model best matching our results is network foraging.     

The evolution of communal roosting in birds: origin and secondary losses

Three main benefits are thought to underlie communal roostingin birds: a reduction in thermoregulation demands, a decreasein predation risk, and an increase in foraging efficiency. Iinvestigated interspecific variation in communal roosting tendenciesacross categories of several ecological factors to examine therelevance of each functional hypothesis in the evolutionary transitionto communal roosting and the secondary reversal to solitary roostinghabits. The study phylogenetic tree included 30 families and437 species. Evolutionary transitions to communal roosting occurredmore often on branches with flocking species and with largerspecies but were not associated with diet, territoriality, geographicalarea, or time of day. The association with flocking activitiessuggests that increased foraging efficiency, a factor thoughtto operate through the formation of flocks, may have been akey factor in the origin of avian communal roosting. However,several transitions to communal roosting occurred on brancheswith nonflocking species, indicating that foraging efficiencymay not be the only factor involved in the evolution of communalroosting. Secondary losses of communal roosting habits occurredon several branches, with a concomitant loss of flocking behaviorand a tendency to exhibit territorial behavior and nocturnalforaging. Secondary losses suggest that communal roosting iscostly to perform and maintain and may be lost when an asocialselection regime operates. The large number of exceptions tothe above patterns may force a reevaluation of current functional hypothesesabout communal roosting in birds.     

ADAPTIVE ADVANTAGES OF MIXED-SPECIES FEEDING FLOCKS AMONG SEED-EATING FINCHES IN COSTA RICA

The feeding ecology of three Costa Rican finches occurring in mixed flocks, Tiaris olivacea, Sporophila aurita corvina and Sporophila torquella , was investigated by measuring both behavioural and ecological variables. Observations on foraging height, rate of hopping and pecking rate, as well as the identity, proximity and number of nearest neighbours were recorded. In addition the duration of all feeding and perching episodes were timed. Comparisons were also made between the abundances of food items (grass seeds) consumed and those potentially available for consumption.
The analysis of variance of the feeding behaviour revealed that the presence or absence of neighbouring birds, whether of the same or different species, influenced the duration of feeding bouts more significantly than did either differences in habitat or species-characteristic behaviour. In addition the dietary comparisons revealed overlap in both species and size of seed consumed. Such similarities suggest that these species are not partitioning fields in the classical sense.
We propose that the increase in the duration of the feeding bout associated with the presence of mixed species aggregations leads to increased feeding efficiency and is the result of intra- and inter-specific social learning. Certainly flocking is often advantageous, since searching in a group facilitates finding clumped resources; mixed species flocking, by increasing exposure to a diversity of foraging places and patterns, can further augment feeding efficiency.     

Does group foraging promote efficient exploitation of resources?

Increased avoidance of food patches previously exploited by other companions has been proposed as one adaptive benefit of group foraging. However, does group foraging really represent the most efficient way to exploit non- or slowly-renewing resources? Here, I used simulations to explore the costs and benefits of exploiting non-renewing resources by foragers searching for food patches independently or in groups in habitats with different types of resource distribution. Group foragers exploited resources in a patch more quickly and therefore spent proportionately more time locating new patches. Reduced avoidance of areas already exploited by others failed to overcome the increased time cost of searching for new food patches and group foragers thus obtained food at a lower rate than solitary foragers. Group foraging provided one advantage in terms of a reduction in the variance of food intake rate. On its own, reduced avoidance of exploitation competition through group foraging appears unlikely to increase mean food intake rate when exploiting non-renewing patches but may provide a way to reduce the risk of an energy shortfall.     

A fine-scale time budget of Cape gannets provides insights into the foraging strategies of coastal seabirds

Central-place foragers organize their feeding trips both to feed themselves and to provide their offspring with food. In seabirds, several long-range foragers have been shown to alternate long and short trips to balance these dual needs. However, the strategies of short-range foragers remain poorly understood. We used a precise, miniaturized motion sensor to examine the time budget of 20 breeding Cape gannets, Morus capensis, foraging off the coast of South Africa. Birds stayed at sea for 5.5-25.3 h, occasionally spending the night at sea. The large number of isolated dives and extended flight time observed during these overnight trips suggested that birds either experienced poor foraging conditions or exploited more distant, yet more profitable prey patches. Conversely, birds that stayed at sea for less than 1 day had relatively consistent activity patterns. Most of these birds (88%) foraged actively at the beginning and at the end of the foraging trip. These feeding bouts were separated by protracted periods of sitting on the sea surface. Such resting periods probably allow birds to digest the food ingested during the first part of the foraging trip, so they initially feed themselves, and then obtain food for their chick on the way back to the breeding site.     

Foraging success in junco flocks and the effects of social hierarchy

The relative advantages of flocking for socially dominant and subordinate dark-eyed juncos (Junco hyemalis) were investigated in a laboratory experiment. Success in finding seeds hidden in an artificial tree was determined for all individuals hunting alone. When hunting alone, birds who spread their searching more evenly over the habitat had a greater chance of finding the food source. Flocks of four were housed together and subsequently each flock formed a dominance hierarchy social structure. High-ranking individuals obtained more seeds when in flocks than when hunting as isolates. In numbers of seeds obtained, socially low-ranking birds gained no advantage in comparison to their performance as isolates. In our experiment, the primary advantage of flocking for low-ranking individuals was the reduced chance of obtaining no food at all.     

The dining etiquette of desert baboons: the roles of social bonds, kinship, and dominance in co-feeding networks

To better understand how individual relationships influence patterns of social foraging in primate groups, we explored networks of co-feeding in wild desert baboons (Papio ursinus). To minimize the risk of aggression and injury associated with contest competition, we expected that individual group members would choose to co-feed with those group-mates that are most likely to show tolerance and a willingness to share food patches. We tested two alternative hypotheses about who those group-mates might be: the "social bonds hypothesis" predicts that preferred foraging partners will be those with whom individuals share strong social bonds, indexed by grooming, whereas the "kinship hypothesis" predicts that preferred foraging partners will be relatives. We also investigated and controlled for the effects of dominance rank, given that competitive ability is known to shape foraging patterns. Social network analyses of over 5,000 foraging events for 14 adults in a single troop revealed that baboon co-feeding was significantly correlated with grooming relationships but not genetic relatedness, and this finding was also true of the female-only co-feeding network. Dominant individuals were also found to be central to the co-feeding network, frequently sharing food patches with multiple group-mates. This polyadic analysis of foraging associations between individuals underlines the importance of dominance and affiliation to patterns of primate social foraging.     

Fine-scale food hoarding decisions in New Zealand Robins ( Petroica australis ): is inter-sexual competition important?

Competition for cache retrieval is hypothesised to influence food hoarding intensity. Previous work has tested this hypothesis by evaluating food hoarding rates during foraging bouts when animals are exposed to different levels of competition for cache retrieval. Little is known about how competition might influence fine-scale food hoarding decisions within foraging bouts. I evaluated fine-scale food hoarding decisions of New Zealand Robins (Petroica australis) by offering mealworms to competitively dominant males and subordinate females, both when they were alone and when they foraged together. I then compared food hoarding rates of sequentially handled prey between sexes and social conditions by assessing how the total number of prey cached increased with the total number of prey handled. Relationships for solitary females, solitary males and paired males were non-linear, indicating that they were more likely to consume initially handled prey, and increasingly likely to cache subsequently handled prey items. Non-linear rates of food hoarding may result from declines in the energetic value of prey that are consumed and stored internally as birds become satiated. Somewhat differently, the relationship for paired females was linear, indicating that paired females make a single food hoarding decision based on bout-level foraging conditions, which results in constant fine-scale food hoarding rates. Constant food hoarding rates in paired females, which experience the strongest competitive effects of any treatment, suggest that food consumption is consistently more advantageous than food hoarding under these conditions, regardless of satiation level. Overall results from this study indicate that New Zealand Robins continuously update food hoarding decisions according to their competitive environment and satiation levels, resulting in scale-dependent patterns in food hoarding intensity.     

Effect of increased food abundance near forest edges on flocking patterns of Coal Tit Parus ater winter groups in mountain coniferous forests

CapsuleIndividuals concentrated near forest edges in bigger social groups than in forest interiors and foraged more on pine cones which were more abundant there.

Aims To evaluate differences in food distribution between forest edges and forest interiors and their effects on the non-breeding flocking patterns of Coal Tit populations inhabiting mountain coniferous forests.

Methods We collected cone production data at forests edges and interiors in mountain pine forests located in the Pyrenees (northeast Iberian peninsula). At the same sites, we also quantified Coal Tit abundance, flocking patterns and foraging behaviour by means of paired bird surveys during autumn and early winter.

Results We recorded a larger abundance of pine cones available on trees along forest edges compared with forest interiors. Coal Tit groups were of bigger size along forest edges, although the number of social groups detected did not differ from forest interiors. Our observations on foraging behaviour supported the hypothesis that differences in flock sizes and overall abundances associated with distance to the edge are due to differences in the availability of pine cones and to the heavier use of these foraging substrates by birds along forest edges.

Conclusions Our results suggest that by changing food distribution, edge effects on pine cone production may be significantly involved in local changes in the social structure of the Coal Tit. An increase in resource heterogeneity and local population density may have important implications at a population level, such as favouring mobility of individuals searching for food resources and thus a transient life, and increasing the costs of territory defence to resident individuals.