A distance-dependent estimation of foraging ranges of neighbouring bird colonies

A reliable estimation of foraging ranges is often an indispensable prerequisite of research in animal ecology and in species conservation. In colonial species, home ranges of members of one colony are frequently overlapping and for practical and theoretical reasons, it can be necessary to assess the foraging areas of whole colonies. Here, we show a method to calculate foraging areas for 13 neighbouring colonies of great white egrets (Ardea alba). Observations of flying individuals and aerial survey data of foraging egrets were combined to in three calculation steps. Firstly, we examined flying paths of egrets from six different colonies to the foraging areas and found an exponential decrease of bird density with increasing distance from the colony. Secondly, we used this distance relationship to calculate the probability of colony membership of 2885 foraging egrets, which were counted by nine aerial surveys. Third, the spatial distribution of these memberships was then basis for calculating the final colony foraging areas. We suggest that distant-dependent foraging patterns could be used in other colonial organisms to assess colony membership in larger sets of survey data. Furthermore, we discuss probable causes for the exponential shape of the distance function in great white egrets.     

Inter- and intra-year variation in foraging areas of breeding kittiwakes (Rissa tridactyla)

While seabird conservation efforts have largely focused on protection from threats at the colony (e.g. reducing disturbance and predation), attention is increasingly being given to implementing protection measures for foraging areas at sea. For this to be effective, important foraging areas must be identified. Although numerous studies have examined seabird foraging behaviour, information is still lacking on the variability in area utilisation within and among breeding seasons. GPS devices were attached to adult black-legged kittiwakes breeding at an expanding North Sea colony (55°20′N, 1°32′W) during both incubation and chick-rearing in 2012 and during chick-rearing in 2011, to determine whether foraging areas remained consistent and to identify the oceanographic characteristics of areas used for foraging. The type and size of prey items consumed at different stages of the breeding cycle was also examined. During incubation (April–May 2012), kittiwakes foraged substantially further from the colony and fed on larger sandeels than when feeding chicks, and there was significant inter-annual variation in foraging areas used during the chick-rearing period (June–July 2011 and 2012). Foraging areas were characterised by cooler sea surface temperatures and areas of high chlorophyll a concentration, although association with specific oceanographic features changed within the breeding season and between years. These results emphasise the importance of considering how foraging areas and reliance on specific oceanographic conditions change over time when seeking to identify important marine areas for seabirds.     

On the integration of individual foraging strategies with colony ergonomics in social insects: nectar-collection in honeybees

Summary We experimentally tested whether foraging strategies of nectar-collecting workers of the honeybee (Apis mellifera) vary with colony state. In particular, we tested the prediction that bees from small, fast growing colonies should adopt higher workloads than those from large, mature colonies. Queenright small colonies were set up by assembling 10 000 worker bees with approximately 4100 brood cells. Queenright large colonies contained 35 000 bees and some 14 500 brood cells. Thus, treatments differed in colony size but not in worker/brood ratios. Differences in workload were tested in the context of single foraging cycles. Individuals could forage on a patch of artificial flowers offering given quantities and qualities of nectar rewards. Workers of small colonies took significantly less nectar in an average foraging excursion (small: 40.1 ± 1.1 SE flowers; large: 44.8 ± 1.1), but spent significantly more time handling a flower (small: 7.3 ± 0.4 s ; large: 5.8 ± 0.4 s). When the energy budgets for an average foraging trip were calculated, individuals from all colonies showed a behavior close to maximization of net energetic efficiency (i.e., the ratio of net energetic gains to energetic costs). However, bees from small colonies, while incurring only marginally smaller costs, gained less net energy per foraging trip than those from large colonies, primarily as a result of prolonged handling times. The differences between treatments were largest during the initial phases of the experimental period when also colony development was maximally different. Our results are at variance with simple models that assume natural selection to have shaped behavior in a single foraging trip only so as to maximize colony growth. Offprint requests to: P. Schmid-Hempel     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

High social density increases foraging and scouting rates and induces polydomy in Temnothorax ants

Many organisms live in crowded groups where social density affects behavior and fitness. Social insects inhabit nests that contain many individuals where physical interactions facilitate information flow and organize collective behaviors such as foraging, colony defense, and nest emigration. Changes in nest space and intranidal crowding can alter social interactions and affect worker behavior. Here, I examined the effects of social density on foraging, scouting, and polydomy behavior in ant colonies—using the species Temnothorax rugatulus. First, I analyzed field colonies and determined that nest area scaled isometrically with colony mass—this indicates that nest area changes proportionally with colony size and suggests that ants actively control intranidal density. Second, laboratory experiments showed that colonies maintained under crowded conditions had greater foraging and scouting activities compared to the same colonies maintained at a lower density. Moreover, crowded colonies were significantly more likely to become polydomous. Polydomous colonies divided evenly based on mass between two nests but distributed fewer, heavier workers and brood to the new nests. Polydomous colonies also showed different foraging and scouting rates compared to the same colonies under monodomous conditions. Combined, the results indicate that social density is an important colony phenotype that affects individual and collective behavior in ants. I discuss the function of social density in affecting communication and the organization of labor in social insects and hypothesize that the collective management of social density is a group level adaptation in social insects.     

Shifting foraging strategies in colonies of the social wasp Polybia occidentalis (Hymenoptera, Vespidae)

Social insect colonies can be expected to forage at rates that maximize colony fitness. Foraging at higher rates would increase the rate of worker production, but decrease adult survival. This trade-off has particular significance during the founding stage, when adults lost are not replaced. Prior work has shown that independent-founding wasps rear the first workers rapidly by foraging at high rates. Foraging rates decrease after those individuals pupate, presumably reducing the risk of foundress death. In the swarm-founding wasps, colony-founding units have many workers, making colony death by forager attrition less likely. Do swarm-founding wasps show similar shifts in foraging rates during the founding stage? We measured foraging rates of the swarm-founding wasp, Polybia occidentalis at four stages of colony development. At each stage, foraging rates correlated with the number of larvae present, which, in the founding stages, correlated with the number of cells in the new nest. Thus, foraging rates appear to be demand-driven, with the level of demand in the founding stage set by the size of nest that is constructed. During the founding stage, foraging rates per larva were high initially, suggesting that colonies minimize the development times of larvae early in the founding stage. Later in the stage, foraging rates decreased, which would reduce worker mortality until new workers eclose. This pattern is similar to that shown for independent-founding wasps and likely results from conflicting pressures to maximize colony growth and minimize the risk of colony death by forager attrition.     

Flexible foraging strategies of gentoo penguins <Emphasis Type="Italic">Pygoscelis papua</Emphasis> over 5 years in the South Shetland Islands,Antarctica

Gentoo penguins Pygoscelis papua show considerable plasticity in their diet, diving, and foraging behaviors among colonies; we expected that they might exhibit similar variability over time, at a single site, since flexible foraging habits would provide a buffer against changes in prey availability. We examined interannual changes in the foraging strategies and diet of gentoo penguins in the South Shetland Islands, Antarctica, over 5 years with variable prey abundance. Antarctic krill Euphausia superba was the primary diet item, and fish the secondary, though the importance of these items varied among years. Diving behavior also varied over time: different dive depth distributions were observed among years. Nonetheless, chick-rearing success remained relatively constant, indicating that gentoo penguins were able to maintain chick provisioning by altering their foraging strategy among years. Variable abundance of krill in the region did not have observable impacts on the diet, foraging behaviors or chick-rearing success of gentoo penguins. We suggest that foraging plasticity may be one reason that gentoo penguin populations have remained stable in the region, while their congeners (P. antarctica and P. adeliae) with less flexible foraging strategies have declined.     

Trade-offs in foraging success and predation risk with spatial position in colonial spiders

Summary Colonial web-building spiders respond to trade-offs between selective forces relative to spatial position within colonies and thus provide support for the selfish herd theory. The size distribution of spiders within colonies of Metepeira incrassata, a colonial orb-weaver (Araneae: Araneidae) from tropical Mexico is nonrandom; larger (mature) spiders and females guarding eggsacs are more prevalent in the center, whereas more small (immature) spiders are found on the periphery. Experimental field studies with spiders of selected size classes show that larger spiders actively and aggressively seek protected positions in the center of the colony webbing, even though prey availability and capture rates are significantly higher on the periphery. Attacks by predatory wasps, other spiders, and hummingbirds are more frequent on the periphery than in the core of the colony. Reproductive females on the periphery are at greater risk because they are captured more often than smaller spiders, and if their egg sacs consequently remain unguarded, chances of cocoon parasitism are increased. As a result, spiders in the core of the colony have greater reproductive success, producing more egg sacs with greater hatching frequency. Colonial spiders thus appear to be making a trade-off between foraging and protection from predation and show a spatial organization predicted by the selfish herd theory. The influence of such trade-offs on individual fitness and the structure of colonies is discussed. Offprint requests to: G.W. Uetz     

Flexible foraging strategies in a diving seabird with high flight cost

How central-place foragers change search strategy in response to environmental conditions is poorly known. Foragers may vary the total distance travelled and how far they range from the central place in response to variation in the distribution of their prey. One potential reason as to why they would extend the length of their foraging trip and its distance from the colony would be to increase prey quality or quantity, despite incurring higher transit costs. To test this trade-off hypothesis in a species with high flight costs, we recorded the foraging behaviour of razorbills (Alca torca) using state-of-the-art techniques that log both individual horizontal (flight activity) and vertical (dive activity) movements. We show that the distance that razorbills travelled to foraging locations increased with sea-surface temperature, which may relate to higher prey quality or quantity. This relation is supported by an indirect index of patch quality, based on dive profiles, which also increased with travel distance from the colony. Furthermore, we show that this index was highest during the daily peak in diving activity, around midday. Taken together, these results suggest that razorbills are capable of adjusting their search strategies sensitively in response to proximate environmental cues.     

Feeding behavior of the sea star Astropecten articulatus (Echinodermata: Asteroidea): an evaluation of energy-efficient foraging in a soft-bottom predator

The energy efficiency of the foraging behavior of Astropecten articulatus (Say) was evaluated in the laboratory. Individuals utilized in the study were collected in the northern Gulf of Mexico from 1990 to 1992. Sea stars presented with equal numbers and weights of low-quality and high-quality prey consistently selected prey of the higher quality. Choice of prey appeared to be mediated by contact chemoreception. Individuals presented with equal weights or equal numbers of different-sized prey models demonstrated a significant preference for smaller prey. Size-selective feeding may be attributable to the ease of manipulation and ingestion of smaller prey, which mazimizes food intake per unit time. In the absence of prey, A. articulatus exhibited a directional pattern of movement. However, as prey were encountered, both the frequency and magnitude of angular deviations in the foraging path increased, resulting in increased foraging in areas of higher prey density. This response to prey availability may increase foraging efficiency by maximizing the rate of prey encounter. Like four other species of the genus Astropecten, A. articulatus exhibited two peaks of activity corresponding with dawn and dusk. Diurnal activities with periods of increased prey availability or periods during which predators are diminished or absent. The sea star A. articulatus exhibits foraging behaviors consistent with the maximization of net energy intake per unit foraging time.     

Effects of prey quantity on predatory wasps (<Emphasis Type="Italic">Polistes dominulus</Emphasis>) when patch quality differs

To determine the effects of prey quantity on central-place foraging of predatory wasps ( Polistes dominulus), prey of varying quality were distributed in patches. A field experiment was conducted, which controlled the amount and quality of prey available. 'Low-fed' colonies were provided with one-third the quantity of prey as that of 'high-fed' colonies. Both were provided with a 1:1 ratio of palatable:unpalatable prey. Experienced wasps of the high-fed colonies never selected unpalatable prey as their first choice of the day, whereas those of the low-fed colonies selected unpalatable prey as their first choice about 1:4 times. In general, wasps from the high-fed colonies reduced palatable patches to zero prey before exploiting unpalatable patches. Foundresses of high-fed colonies captured disproportionately more palatable prey than those of low-fed nests, but there was no correlation between ratio of palatable to unpalatable prey taken and the number of offspring produced. Wasps from low-fed colonies attacked unpalatable prey sooner, but not without considerable effort to avoid use of those patches. Foundresses of low-fed colonies also spent a greater proportion of time overall in unsuccessful search, which may explain why only wasps from low-fed colonies foraged on cool days. High-fed colonies produced more cells and more and heavier offspring than low-fed colonies. But the productivity of the low-fed colonies was greater than that of the 'natural' colonies, which had to find their own prey in a field-woodland area. These results indicate that prey scarcity changes foraging behavior and affects prey choice. These changes may not totally alleviate negative effects of unpalatable prey on colony development and offspring production. The results of this study increase understanding of the central-place foraging behavior of paper wasps, which are important biocontrol agents in natural and agricultural settings.     

Relatedness structure and kin-biased foraging in the greater horseshoe bat (Rhinolophus ferrumequinum)

Female greater horseshoe bats (Rhinolophus ferrumequinum) exhibit strong natal philopatry to their maternity roost over many years, leading to the aggregation of matrilineal kin. Maternity colonies may, therefore, be expected to comprise highly related individuals, and, as such, provide conditions suitable for the evolution of kin-selected behaviours. To test these predictions, we examined relatedness and behaviour among matrilineal kin within a colony in south-west Britain. Genetic analysis of 15 matrilines, identified from microsatellite genotyping and long-term ringing surveys, revealed average relatedness levels of 0.17 to 0.64. In contrast, background relatedness among colony females approximated to zero (0.03). These results suggest that inclusive fitness benefits may only be accrued through discriminate cooperation within matrilines, and not at the wider colony level. To examine whether the potential for such benefits is realised through kin- biased cooperation during foraging, females from two matrilines were radio-tracked simultaneously over 3 years. Pairwise home-range overlap correlated significantly with Hamilton's relatedness coefficient. The greatest spatial associations were observed between females and their adult daughters, which shared both foraging grounds and night roosts, sometimes over several years. Tagged females, however, generally foraged and roosted alone, suggesting that kin-biased spatial association probably does not result from either information-transfer or cooperative territorial defence. Such patterns may instead result from a mechanism of maternal inheritance of preferred foraging and roosting sites.     

How resources and encounters affect the distribution of foraging activity in a seed-harvesting ant

We examined how the foraging ecology of the seed-harvesting ant Messor andrei depends upon the distribution of resources and the presence of conspecifics. Bait experiments showed that colonies can recruit to high-density patches of seeds. However, at the seasonal scale, natural resource distribution did not affect the distribution of foraging activity. We conducted the study in years of high rainfall and thus seed availability may not have been a limiting factor. Colonies always preferred to forage in areas closer to their nest, which may reduce travel time between the nest and foraging sites. On a day-to-day scale, encounters between neighboring colonies at a site increased the probability that colonies would return to forage at that site; this was true both for natural and experimental encounters. In the summer, this resulted in colonies foraging at the sites of intraspecific encounters on more days than in areas where no encounter had occurred. Encounters between colonies included fighting, and there was little overlap between the foraging areas of neighboring colonies: both results suggest that one function of encounters is to defend foraging space. The high probability of return to the site of an encounter between colonies suggests that encounters may have a second function: to indicate the presence of resources. Received: 28 June 1999 / Received in revised form: 12 October 1999 / Accepted: 16 October 1999     

Parasitism in a social wasp: effect of gregarines on foraging behavior, colony productivity, and adult mortality

Behavior in eusocial insects likely reflects a long history of selection imposed by parasites and pathogens because the conditions of group living often favor the transmission of infection among nestmates. Yet, relatively few studies have quantified the effects of parasites on both the level of individual colony members and of colony success, making it difficult to assess the relative importance of different parasites to the behavioral ecology of their social insect hosts. Colonies of Polybia occidentalis, a Neotropical social wasp, are commonly infected by gregarines (Phylum Apicomplexa; Order Eugregarinida) during the wet season in Guanacaste, Costa Rica. To determine the effect of gregarine infection on individual workers in P. occidentalis, we measured foraging rates of marked wasps from colonies comprising both infected and uninfected individuals. To assess the effect of gregarines on colony success, we measured productivity and adult mortality rates in colonies with different levels of infection prevalence (proportion of adults infected). Foraging rates in marked individuals were negatively correlated with the intensity of gregarine infection. Infected colonies with high gregarine prevalence constructed nests with fewer brood cells per capita, produced less brood biomass per capita, and, surprisingly, experienced lower adult mortality rates than did uninfected or lightly infected colonies. These data strongly suggest that gregarine infection lowers foraging rates, thus reducing risk to foragers and, consequently, reducing adult mortality rates, while at the same time lowering per-capita input of materials and colony productivity. In infected colonies, queen populations were infected with a lower prevalence than were workers. Intra-colony infection prevalence decreased dramatically in the P. occidentalis population during the wet season.An erratum to this article can be found at     

Effects of interactions among genotypically diverse nestmates on task specialization by foraging honey bees (Apis mellifera)

Summary Recent studies have shown that differences in patterns of task specialization among nestmate honeybee workers (Apis mellifera) can be explained, in part, as a consequence of genotypic variability. Here, we present evidence supporting the hypothesis that an individual's pattern of task specialization is affected not only by her own genotype, but, indirectly, by the genotypes of her nestmates. Workers from two strains of honey bees, one selected for high pollen hoarding, the other for low pollen hoarding, were observed in colonies of their respective parent strains and in colonies of the other strain. Worker genotype and host-colony type affected foraging activity. Workers from the high strain fostered in low-strain colonies returned with pollen on 75.6% of total foraging trips, while workers from the high strain fostered in high-strain colonies returned with pollen on 53.5% of total trips. Workers from the low strain fostered in low-strain colonies returned with pollen on 34.8% of total foraging trips while workers from the low strain fostered in high-strain colonies returned with pollen on 2.6% of total trips. Similar results were obtained in a second experiment. We suggest that workers influence the behavior of their nestmates indirectly through their effects on the shared colony environment. The asymmetry seen in the response of workers from these strains to the two types of colony environments also suggests that these genotypes exhibit different norms of reaction. Offprint requests to: N.W. Calderone     

How colony growth affects forager intrusion between neighboring harvester ant colonies

Summary Colonies of the harvester ant, Pogonomyrmex barbatus, adjust the direction and length of foraging trails in response to the foraging behavior of their conspecific neighbors. In the absence of any interaction with its neighbor, a mature colony expands its foraging range at a rate of 0.85 ± 0.15 m per day. Exclusion experiments show that if a colony is prevented from using its foraging trails, the neighbors of that colony will enter its foraging range within 10 days. Exclusion experiments were performed with three age classes of colonies: young (1 year old), intermediate (3–4 years old), and old (5 years old or more). Colonies 3–4 years old are most likely to expand foraging ranges, and to retain newly-gained areas. To examine the relation of colony age (in years) and colony size (in numbers of workers), colonies of known age were excavated. Colonies increase greatly in size in years 3 and 4. Foraging area may be of greater current or prospective value for younger, smaller, quickly growing colonies than for older, larger ones of stable size. Correspondence to the second address     

How age and sex drive the foraging behaviour in the king penguin

As predicted by life history theory, once recruited into the breeding population and with increasing age, long-lived animals should be able to manage more efficiently the conflict between self-maintenance and reproduction. Consequently, breeding performances should improve with age before stabilizing at a certain level. Using temperature–depth recorders and isotopic analysis, we tested how age affects the foraging behaviour of king penguin Aptenodytes patagonicus during one trip in the chick-rearing phase. Depending on sex and age, king penguins expressed two different foraging strategies. Older birds gained more daily mass per unit body mass than younger ones. Older females conducted shorter trips, dived deeper and performed more prey pursuits. They also had higher blood levels of δ¹⁵N than younger individuals and males indicating sex- and age-specific dietary regimes. However, we found no differences in carbon isotopic signature, suggesting that individuals exploited the same foraging areas independently of sex and age. Our results suggest that king penguins are able to increase the quantity of energy extracted with increasing age and that such a strategy is sex-related. Our study is the first to reveal of an interaction between age and sex in determining foraging efficiency in king penguins.     

Flexible foraging movements of leatherback turtles across the North Atlantic Ocean

Some marine species have been shown to target foraging at particular hotspots of high prey abundance. However, we show here that in the year after a nesting season, female leatherback turtles (Dermochelys coriacea) in the Atlantic generally spend relatively little time in fixed hotspots, especially those with a surface signature revealed in satellite imagery, but rather tend to have a pattern of near continuous traveling. Associated with this traveling, distinct changes in dive behavior indicate that turtles constantly fine tune their foraging behavior and diel activity patterns in association with local conditions. Switches between nocturnal vs. diurnal activity are rare in the animal kingdom but may be essential for survival on a diet of gelatinous zooplankton where patches of high prey availability are rare. These results indicate that in their first year after nesting, leatherback turtles do not fit the general model of migration where responses to resources are suppressed during transit. However, their behavior may be different in their sabbatical years away from nesting beaches. Our results highlight the importance of whole-ocean fishing gear regulations to minimize turtle bycatch.     

Maximization of foraging efficiency and resource defense by group retrieval in the ant Formica schaufussi

Summary Workers of the ant Formica schaufussi forage as individuals and cooperate in groups to retrieve arthropod prey. In 2 sample years, group-transported prey were on average 6.8 and 4.7 times heavier than individually retrieved items, and the average loading ratios of groups were greater than the loading ratios of single foragers. Retrieval group size was adjusted to prey size, and prey transport velocity for individuals and groups tended to decrease with increasing prey weight. The efficiency of individual and group retrieval, estimated from calculations of the prey delivery rate to the nest (PDR) achieved by each foraging mode, varied as a function of prey size. Individual retrieval maximized PDR at a prey weight of 19.5 mg, and group transport maximized PDR at 190 mg. Although the PDR maxima of an individual in a group and a solitary forager were approximately equal, depending on prey size, group transport may maximize foraging efficiency. Group transport also decreased interference competition from sympatric ant species. Group-transported prey having a greater likelihood of successful retrieval were within the size range of prey that maximized foraging efficiency. Transport group size appeared to be more important in prey defense than in increasing prey transport velocity, suggesting an important role of group size in competitive ability.Offprint requests to: J.F.A. Traniello