Feast or famine: evidence for mixed capital–income breeding strategies in Weddell seals

Evolved patterns of resource expenditure for reproduction have resulted in a life history continuum across species. A strictly capital-breeding strategy relies extensively on stored energy for reproduction, whereas income breeding uses energy acquired throughout the reproductive period. However, facultative income breeding has been shown in some classically capital-breeding animals, and was originally thought to provide a nutritional refuge for smaller females incapable of securing sufficient reserves during pre-partum foraging. We examined milk composition and milk output for the Weddell seal to determine to what degree lactation was aided by food intake, and what factors contributed to its manifestation. Milk composition was independent of maternal post-partum mass and condition, but did change over lactation. Changes were most likely in response to energetic and nutritional demands of the pup at different stages of development. During early lactation, females fasted and devoted 54.9% of total energy loss to milk production. Later in lactation 30.5% more energy was devoted to milk production and evidence suggested that larger females fed more during lactation than smaller females. It appears that Weddell seals may exhibit a flexible strategy to adjust reproductive investment to local resource levels by taking advantage of periods when prey are occasionally abundant, although it is restricted to larger females possessing the physiological capacity to dive for longer and exploit different resources during lactation. This supports the assumption that although body mass and phylogenetic history explain most of the variation in lactation patterns (20–69%), the remaining variation has likely resulted from physiological adaptations to local environmental conditions. Our study confirms that Weddell seals use a mixed capital–income breeding strategy, and that considerable intraspecific variation exists. Questions remain as to the amount of energy gain derived from the income strategy, and the consequences for pup condition and survival. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Prospecting and informed dispersal: Understanding and predicting their joint eco‐evolutionary dynamics

The ability of individuals to leave a current breeding area and select a future one is important, because such decisions can have multiple consequences for individual fitness, but also for metapopulation dynamics, structure, and long‐term persistence through non‐random dispersal patterns. In the wild, many colonial and territorial animal species display informed dispersal strategies, where individuals use information, such as conspecific breeding success gathered during prospecting, to decide whether and where to disperse. Understanding informed dispersal strategies is essential for relating individual behavior to subsequent movements and then determining how emigration and settlement decisions affect individual fitness and demography. Although numerous theoretical studies have explored the eco‐evolutionary dynamics of dispersal, very few have integrated prospecting and public information use in both emigration and settlement phases. Here, we develop an individual‐based model that fills this gap and use it to explore the eco‐evolutionary dynamics of informed dispersal. In a first experiment, in which only prospecting evolves, we demonstrate that selection always favors informed dispersal based on a low number of prospected patches relative to random dispersal or fully informed dispersal, except when individuals fail to discriminate better patches from worse ones. In a second experiment, which allows the concomitant evolution of both emigration probability and prospecting, we show the same prospecting strategy evolving. However, a plastic emigration strategy evolves, where individuals that breed successfully are always philopatric, while failed breeders are more likely to emigrate, especially when conspecific breeding success is low. Embedding information use and prospecting behavior in eco‐evolutionary models will provide new fundamental understanding of informed dispersal and its consequences for spatial population dynamics.     

When to start and when to stop: Effects of climate on breeding in a multi‐brooded songbird

Climate warming has been shown to affect the timing of the onset of breeding of many bird species across the world. However, for multi‐brooded species, climate may also affect the timing of the end of the breeding season, and hence also its duration, and these effects may have consequences for fitness. We used 28 years of field data to investigate the links between climate, timing of breeding, and breeding success in a cooperatively breeding passerine, the superb fairy‐wren (Malurus cyaneus). This multi‐brooded species from southeastern Australia has a long breeding season and high variation in phenology between individuals. By applying a “sliding window” approach, we found that higher minimum temperatures in early spring resulted in an earlier start and a longer duration of breeding, whereas less rainfall and more heatwaves (days > 29°C) in late summer resulted in an earlier end and a shorter duration of breeding. Using a hurdle model analysis, we found that earlier start dates did not predict whether or not females produced any young in a season. However, for successful females who produced at least one young, earlier start dates were associated with higher numbers of young produced in a season. Earlier end dates were associated with a higher probability of producing at least one young, presumably because unsuccessful females kept trying when others had ceased. Despite larger scale trends in climate, climate variables in the windows relevant to this species’ phenology did not change across years, and there were no temporal trends in phenology during our study period. Our results illustrate a scenario in which higher temperatures advanced both start and end dates of individuals’ breeding seasons, but did not generate an overall temporal shift in breeding times. They also suggest that the complexity of selection pressures on breeding phenology in multi‐brooded species may have been underestimated.     

Interspecific variation in the phenology of advertisement calling in a temperate Australian frog community

Spatial and temporal partitioning of resources underlies the coexistence of species with similar niches. In communities of frogs and toads, the phenology of advertisement calling provides insights into temporal partitioning of reproductive effort and its implications for community dynamics. This study assessed the phenology of advertisement calling in an anuran community from Melbourne, in southern Australia. We collated data from 1432 surveys of 253 sites and used logistic regression to quantify seasonality in the nightly probability of calling and the influence of meteorological variables on this probability for six species of frogs. We found limited overlap in the predicted seasonal peaks of calling among these species. Those shown to have overlapping calling peaks are unlikely to be in direct competition, due to differences in larval ecology (Crinia signifera and Litoria ewingii) or differences in calling behavior and acoustics (Limnodynastes dumerilii and Litoria raniformis). In contrast, closely related and ecologically similar species (Crinia signfera and Crinia parinsignifera; Litoria ewingii and Litoria verreauxii) appear to have staggered seasonal peaks of calling. In combination with interspecific variation in the meteorological correlates of calling, these results may be indicative of temporal partitioning of reproductive activity to facilitate coexistence, as has been reported for tropical and temperate anurans from other parts of the globe.     

High foraging site fidelity and spatial segregation among individual great black‐backed gulls

Individual foraging site fidelity, whereby individuals repeatedly visit the same foraging areas, is widespread in nature, and likely benefits individuals through higher foraging efficiency and potentially, higher breeding success. It may arise as a consequence of habitat or resource specialisation, or alternatively, where resources are abundant or predictable, the partitioning of space might guarantee individuals exclusive foraging opportunities. We tracked seven adult great black‐backed gulls Larus marinus at a North Sea colony from early incubation to the end of the breeding season in 2016, providing a total of 1170 foraging trips over a mean ± SD tracking period of 67 ± 16 days. There was clear spatial segregation between individuals, with almost no overlap of their core areas (50% utilisation distribution) during incubation and chick‐rearing. Core areas were relatively small and there was high repeatability (R ± SE) in foraging parameters, including initial departure direction (0.73 ± 0.11), foraging range (0.41 ± 0.14) and cumulative distance travelled (0.19 ± 0.1) throughout the breeding season. Despite the low spatial overlap, there was little evidence of differential habitat use by individuals. The near‐exclusive individual foraging areas of this species, usually considered to be a generalist, indicate that where there is high resource availability throughout the breeding season and a small local population, individuals appear to adopt a territorial strategy which likely reduces intraspecific competition.     

State dependence,personality, and plants: light‐foraging decisions in Mimosa pudica (L.)

Plants make foraging decisions that are dependent on ecological conditions, such as resource availability and distribution. Despite the field of plant behavioral ecology gaining momentum, ecologists still know little about what factors impact plant behavior, especially light‐foraging behavior. We made use of the behavioral reaction norm approach to investigate light foraging in a plant species that exhibits rapid movement: Mimosa pudica. We explored how herbivore avoidance behavior in M. pudica (which closes its leaflets temporarily when disturbed) is affected by an individual's energy state and the quality of the current environment and also repeatedly tested the behavior of individuals from two seed sources to determine whether individuals exhibit a “personality” (i.e., behavioral syndrome). We found that when individuals are in a low‐energy state, they adopt a riskier light‐foraging strategy, opening leaflets faster, and not closing leaflets as often in response to a disturbance. However, when plants are in a high‐energy state, they exhibit a plastic light‐foraging strategy dependent on environment quality. Although we found no evidence that individuals exhibit behavioral syndromes, we found that individuals from different seed sources consistently behave differently from each other. Our results suggest that plants are capable of making state‐dependent decisions and that plant decision making is complex, depending on the interplay between internal and external factors.     

Reproductive biology of Melanophryniscus montevidensis (Anura: Bufonidae) from Uruguay: reproductive effort,fecundity, sex ratio and sexual size dimorphism

We analyzed the reproductive traits of a population of Melanophryniscus montevidensis. The reproductive events were associated to rainfalls. Males called from dawn to late night, were more abundant than females, and adopted alternative tactics to find partners (active search). Females were larger than males, and had mature oocytes and lipid reserves after spawning; they showed no association between size and fecundity. As an explosive breeder, females are prepared all year long for reproduction. Several individuals revealed high site-fidelity for breeding. Knowledge about breeding pattern could help to propose suitable and efficient in situ conservation measures for this endangered species.     

Use of Social and Ecological Information in Tamarin Foraging Decisions

A major consequence of group living is that foragers may rely on social information in addition to ecological information to locate feeding sites. Although conspecifics can provide cues as to the spatial location of food patches, individual foraging decisions also must include some assessment of the likelihood of obtaining access to a resource other group members seek. This likelihood differs in the 2 models generally proposed to explain intragroup social foraging: the information-sharing model and the producer-scrounger model. We conducted an experimental field study on wild groups of emperor (Saguinus imperator) and saddleback (S. fuscicollis) tamarins to determine the foraging strategies adopted by individual group members and their relationship to social rank, food intake, and the ability to use ecological and social information in making intra-patch foraging decisions. Individual tamarins applied different behavioral strategies compatible with a finder-joiner paradigm to solve foraging problems. About half of the individuals in each study group initiated 74%–90% of all food searches and acted as finders. Most alpha individuals adopted a joiner strategy by monitoring the activities of others' to obtain a reward. The individual arriving first at a reward platform enjoyed a finder's advantage. Despite differences in search effort, both finders and joiners presented similar abilities in learning to associate ecological cues with the presence of food rewards at our experimental feeding stations. We conclude that within a group foraging context, tamarins integrate social and ecological information in decision-making.     

Changes in breeding phenology of eastern Ontario frogs over four decades

Global climate change has been implicated in phenological shifts for a variety of taxa. Amphibian species in particular are sensitive to changes in their environment due to their biphasic life history and restricted reproductive requirements. Previous research has shown that not all temperate amphibian species respond similarly to the same suite of climatic or environmental cues, nor are individual species necessarily uniform in their responses across their range. We examined both the timing of spring emergence and calling phenology of eight anuran species in southeastern Ontario, Canada, using an approximately 40‐year dataset of historical records of amphibian activity. Rana pipiens was the only species out of eight considered to emerge significantly earlier, by an estimated 22 days over four decades. Both R. pipiens and Bufo americanus have advanced initiation of calling over a four‐decade span significantly earlier by an estimated 37.2 and 19.2 days, respectively. Rana sylvatica showed a trend toward earlier emergence by 19 days, whereas we did not detect changes in emergence phenology for the remaining five species. This significant shift in breeding behavior for two species correlates to significant regional increases in spring temperatures of an estimated 2.7–2.8°C overall over four decades. Our study suggests that local temperature increases have affected the timing of emergence and the onset of calling activity in some Ontario anuran species. Global decline or range shifts ultimately may be related to changes in reproductive behavior and timing mediated by shifting climate.     

Steroid hormone profiles and relative body condition of calling and satellite toads: implications for proximate regulation of behavior in anurans

Males of most anuran species (frogs and toads) vocalize to attractmates. However, individuals of many vocal species may also adoptalternative noncalling "satellite" tactics. Satellite malescharacteristically remain in close proximity to calling conspecificsand attempt to intercept incoming females attracted to advertisingmales. Emerson proposed that alternation between calling andnoncalling behavior in anurans is mediated by a reciprocal interactionbetween circulating levels of corticosterone and androgens thatis driven by depletion of energy reserves during vocalization.We tested this hypothesis by examining steroid hormone profilesand the relative body condition of calling and satellite Woodhouse'stoads (Bufo woodhousii) and Great Plains toads (B. cognatus).Consistent with Emerson's hypothesis, callers had significantlyhigher circulating corticosterone levels and were in bettercondition than satellites. However, levels of testosterone anddihydrotestosterone did not differ significantly between satellitesand callers, and we found no evidence that high levels of corticosteronehad an inhibitory effect on androgen production in either species.These data thus support a relationship between corticosteronelevels and depletion of energy reserves during bouts of vocalizationbut suggest that alternation between calling and satellite behaviormay be associated with direct effects of corticosterone on brainvocal control centers. We propose a model that incorporatesrelationships among energy reserves, androgens, corticosterone,and arginine vasotocin-producing neurons in the telencephalonto explain transitions between calling and satellite tacticsin toads.     

Plasma mammalian leptin analogue predicts reproductive phenology,but not reproductive output in a capital‐income breeding seaduck

To invest in energetically demanding life history stages, individuals require a substantial amount of resources. Physiological traits, particularly those related to energetics, can be useful for examining variation in life history decisions and trade‐offs because they result from individual responses to environmental variation. Leptin is a protein hormone found in mammals that is proportional to the amount of endogenous fat stores within an individual. Recently, researchers have confirmed that a mammalian leptin analogue (MLA), based on the mammalian sequence of leptin, is present with associated receptors and proteins in avian species, with an inhibitory effect on foraging and body mass gain at high circulating levels. While MLA has been both quantified and manipulated in avian species, little is currently known regarding whether plasma MLA in wild‐living species and individuals is associated with key reproductive decisions. We quantified plasma MLA in wild, Arctic‐nesting female common eiders (Somateria mollissima) at arrival on the breeding grounds and followed them to determine subsequent breeding propensity, and reproductive phenology, investment, and success. Common eiders are capital‐income breeding birds that require the accumulation of substantial fat stores to initiate laying and successfully complete incubation. We found that females with lower plasma MLA initiated breeding earlier and in a shorter period of time. However, we found no links between plasma MLA levels and breeding propensity, clutch size, or reproductive success. Although little is still known about plasma MLA, based on these results and its role in influencing foraging behaviors and condition gain, plasma MLA appears to be closely linked to reproductive timing and is therefore likely to underlie trade‐offs surrounding life history decisions.     

Foraging strategies of the acellular slime moulds Didymium iridis and Didymium bahiense

Organisms can increase their foraging efficiency by modifying their behaviour according to information about the quality of currently exploited resource patches. Here we examine the effect of food concentration on the foraging strategies of two previously unstudied species of slime mould: Didymium iridis and Didymium bahiense. We studied two main foraging decisions: how long to wait before commencing exploration of the surrounding environment (exploitation strategy) and how intensely to search the environment for new opportunities (exploration strategy). Food concentration did not affect exploitation behaviour in either D. iridis or D. bahiense. Food concentration did affect exploration behaviour in D. iridis, but not in D. bahiense. Encounters with food resources, irrespective of concentration, resulted in increased exploitation and decreased exploration in D. iridis but did not influence foraging behaviour in D. bahiense. We suggest that the varying foraging strategies of slime moulds may have evolved to exploit different resource distributions in their natural environments. We also discuss the potential impact of microbial contamination and differences in handling regimes.     

Optimal parasite infection strategies: a state-dependent approach

Many macroparasites spend a crucial phase of their life-cycle as free-living infective stages. Despite their importance, however, little theoretical work has considered how evolution may shape the behaviour of these stages. Here, we develop what we believe to be the first stochastic dynamic programming model of parasite life-history strategies to investigate how a trade-off between resource depletion and host encounter rate may shape the optimal infection strategy of a macroparasite. The optimal strategy depends strongly on the probability of host contact and, depending on the relative costs and benefits, macroparasites should adopt either a passive 'ambushing' (sit-and-wait) strategy, an active 'cruising' strategy or a mixed strategy with an initial cruising phase, followed by a switch to ambushing when energy reserves fall to a threshold level. Under no circumstances does the model predict ambush-then-cruise. We use our model to help interpret previously published data on entomopathogenic nematode foraging behaviour, showing how this framework could facilitate our understanding of macroparasite behaviour during this key stage of the life-cycle.     

Predation-Related Costs and Benefits of Conspecific Attraction in Songbirds—An Agent-Based Approach

Songbirds that follow a conspecific attraction strategy in the habitat selection process prefer to settle in habitat patches already occupied by other individuals. This largely affects the patterns of their spatio-temporal distribution and leads to clustered breeding. Although making informed settlement decisions is expected to be beneficial for individuals, such territory clusters may potentially provide additional fitness benefits (e.g., through the dilution effect) or costs (e.g., possibly facilitating nest localization if predators respond functionally to prey distribution). Thus, we hypothesized that the fitness consequences of following a conspecific attraction strategy may largely depend on the composition of the predator community. We developed an agent-based model in which we simulated the settling behavior of birds that use a conspecific attraction strategy and breed in a multi-predator landscape with predators that exhibited different foraging strategies. Moreover, we investigated whether Bayesian updating of prior settlement decisions according to the perceived predation risk may improve the fitness of birds that rely on conspecific cues. Our results provide evidence that the fitness consequences of conspecific attraction are predation-related. We found that in landscapes dominated by predators able to respond functionally to prey distribution, clustered breeding led to fitness costs. However, this cost could be reduced if birds performed Bayesian updating of prior settlement decisions and perceived nesting with too many neighbors as a threat. Our results did not support the hypothesis that in landscapes dominated by incidental predators, clustered breeding as a byproduct of conspecific attraction provides fitness benefits through the dilution effect. We suggest that this may be due to the spatial scale of songbirds’ aggregative behavior. In general, we provide evidence that when considering the fitness consequences of conspecific attraction for songbirds, one should expect a trade-off between the benefits of making informed decisions and the costs of clustering.     

Alternative reproductive tactics in snail shell‐brooding cichlids diverge in energy reserve allocation

Life history theory predicts that the amount of resources allocated to reproduction should maximize an individual's lifetime reproductive success. So far, resource allocation in reproduction has been studied mainly in females. Intraspecific variation of endogenous energy storage and utilization patterns of males has received little attention, although these patterns may vary greatly between individuals pursuing alternative reproductive tactics (ARTs). ARTs are characterized by systematic variation of behavioral, physiological, and often morphological traits among same‐sex conspecifics. Some individuals may rely on previously accumulated reserves, because of limited foraging opportunities during reproduction. Others may be able to continue foraging during reproduction, thus relying on reserves to a lesser extent. We therefore predicted that, if male tactics involve such divergent limitations and trade‐offs within a species, ARTs should correspondingly differ in energy reserve allocation and utilization. To test this prediction, we studied short‐term and long‐term reserve storage patterns of males in the shell‐brooding cichlid Lamprologus callipterus. In this species, bourgeois males investing in territory defense, courtship, and guarding of broods coexist with two distinct parasitic male tactics: (1) opportunistic sneaker males attempting to fertilize eggs by releasing sperm into the shell opening when a female is spawning; and (2) specialized dwarf males attempting to enter the shell past the spawning female to fertilize eggs from inside the shell. Sneaker males differed from other male types by showing the highest amount of accumulated short‐term and long‐term fat stores, apparently anticipating their upcoming adoption of the nest male status. In contrast, nest males depleted previously accumulated energy reserves with increasing nest holding period, as they invest heavily into costly reproductive behaviors while not taking up any food. This conforms to a capital breeder strategy. Dwarf males did not accumulate long‐term fat stores at all, which they can afford due to their small behavioral effort during reproduction and their continued feeding activity, conforming to an income breeder strategy. Our data confirm that the resource storage patterns of males pursuing ARTs can diverge substantially, which adds to our understanding of the coexistence and maintenance of alternative reproductive patterns within species.     

Potential Component Allee Effects and Their Impact on Wetland Management in the Conservation of Endangered Anurans

Effective management of wetland quantity and quality is crucial for effective conservation of declining amphibian populations. In particular, frogs and toads that employ aggregative breeding strategies may suffer negative population impacts in response to changes in availability of aquatic breeding habitat, including overabundance of suitable habitat, if density of conspecifics attending aggregations is positively correlated with reproductive success. Here we document such a positive relationship, potentially the first example of a component Allee effect in an anuran, in the critically endangered Houston toad (Bufo houstonensis). We assessed the relationship between mean yearly chorus size and reproductive success of males at the pond level using an information theoretic model selection approach and a two-sample t-test. The chosen model contained the single variable of mean yearly chorus size to predict probability of reproduction, as selected using the Akaike Information Criterion corrected for small sample size and Akaike weight. Mean chorus sizes were significantly higher among ponds exhibiting evidence of reproduction than in those that showed no evidence of reproduction. Our results suggest that chorusing alone is a poor proxy for inference of population stability and highlight a need for reassessment of widely-used amphibian monitoring protocols. Further, amphibian conservation efforts should account for potential Allee effects in order to optimize benefits and avoid underestimating critical population thresholds, particularly in species exhibiting rapid population declines.     

Adjustment of pre‐moult foraging strategies in Macaroni Penguins Eudyptes chrysolophus according to locality,sex and breeding status

The annual moult creates the highest physiological stress during a penguin's breeding‐cycle and is preceded by a period of hyperphagia at sea. Although crucial to individual survival, foraging strategies before moult have been little investigated in keystone marine consumers in the Southern Ocean. The Macaroni Penguin Eudyptes chrysolophus demonstrates how individuals may adjust their foraging strategies during this period in line with constraints such as potential intraspecific competition between localities, foraging ability between dimorphic sexes and timing at sea between breeding and non‐breeding population components. We recorded pre‐moult behaviour at sea for 22 Macaroni Penguins from Crozet and Kerguelen Islands (southern Indian Ocean) during 2009 and 2011, using light‐based geolocation and stable isotope analysis. Penguins were distributed in population‐specific oceanic areas with similar surface temperatures (3.5 °C) south of the archipelagos, where they foraged at comparable trophic levels based on stable isotopes of their blood. Bayesian ‘broken stick’ modelling with concurrent analysis of seawater temperature records from the animal‐borne devices showed that within each population, females remained 6 days longer than males in the colder waters before heading back towards their colonies. Finally, 17 other non‐breeding individuals that moulted earlier had a higher mean blood δ¹⁵N value than did post‐breeding birds, meaning that early moulters probably fed more on fish than did late moulters. Our findings of such adjustments in foraging strategies developed across locality, sex and breeding status help understanding of the species' contrasted pre‐moult biology across its range and its ecology in the non‐breeding period.     

Chorusing Behaviour,a Density-dependent Alternative Mating Strategy in Male Common Toads (Bufo bufo)

Male chorusing behaviour was studied in a population of common toads (Bufo bufo) on the island of Öland south Sweden, and the functional role of male advertisement calling in this species was experimentally examined. Calling males were larger and heavier than non-calling males (t = 2.41, p < 0.025 and t = 2.36, p < 0.025, respectively). However, small males were also found to call. This indicates that large males persisted in calling for longer and/or called more often. The proportion of calling males decreased as population size increased during the breeding season, indicating that calling is a low density strategy. Females responded more readily to calls than males. There were insufficient data to determine if the dominant frequencies of advertisement calls were inversely correlated with male body size, however, this relationship was found for the similar release calls. Females were found not to discriminate between high and low frequency calls, but when given a choice between two calls of different sound pressure levels (SPL), females were attracted to the louder calls. Thus, the function of chorusing is to advertise the position of males to mate-seeking females when the population density is low.     

Calling phenology and detectability of a threatened amphibian (Litoria olongburensis) in ephemeral wetlands varies along a latitudinal cline: Implications for management

Amphibian reproductive biology and activity are highly sensitive to environmental cues. For amphibian conservation in the face of climate change, a clear priority is to investigate impacts of variable weather on breeding activity and detectability. This study assessed variation in the phenology, intensity and detectability of calling for a threatened temporary wetland breeding species, Litoria olongburensis. Acoustic and visual surveys were conducted monthly from August 2009 to July 2011 over the species latitudinal range in eastern Australia. Calling phenology of L. olongburensis varied spatially and temporally within and among latitudinal clusters. Detectability of frogs also varied among months and transects, and also with the type of survey conducted (visual vs. acoustic). Air temperature was a strong predictor for the onset of the calling season (early spring); however, peaks in calling abundances were primarily driven by monthly accumulated rainfall. The duration of the calling season varied among latitudinal clusters, becoming narrower moving south, reflecting clinal variation in weather patterns. This study presents a framework for amphibian species management using a combination of survey techniques and the climatic conditions influencing detectability. Applied along the latitudinal gradient, it provides data for predicting the likely impacts of climate change on amphibian activity and reproduction.     

The role of information transfer under different food patterns: a simulation study

The aim of this paper is to address the question of how informationtransfer affects foraging efficiency in a colonial breedingsituation. By creating computer simulations, we attempt to modelinformation transfer of food location by individuals withinthe colony. Three kinds of foraging strategy were modeled: searcher–noinformation transfer (solo foraging), watcher–limitedinformation transfer (local enhancement), and a mixture of watcherand follower–full information transfer (information center).The predictability of food was changed by varying patchiness(ratio of food patches) and duration of food patches. When thefood occurs in many randomly distributed patches and the fooddensity in each is low, the solitary searcher strategy givesthe best foraging efficiency. The significance of informationtransfer strategies and colonial breeding on foraging efficiencyincrease when food becomes more clumped. The solitary watcherstrategy is the best at intermediately clumped food distribution.Colonial breeding gives the best foraging efficiency when theinformation center operates and there are some high-densityfood patches.