Abstract: Network analysis has a long history in the mathematical and social sciences and the aim of this introduction is to provide a brief overview of the potential that it holds for the study of animal behaviour. One of the most attractive features of the network paradigm is that it provides a single conceptual framework with which we can study the social organisation of animals at all levels (individual, dyad, group, population) and for all types of interaction (aggressive, cooperative, sexual etc.). Graphical tools allow a visual inspection of networks which often helps inspire ideas for testable hypotheses. Network analysis itself provides a multitude of novel statistical tools that can be used to characterise social patterns in animal populations. Among the important insights that networks have facilitated is that indirect social connections matter. Interactions between individuals generate a social environment at the population level which in turn selects for behavioural strategies at the individual level. A social network is often a perfect means by which to represent heterogeneous relationships in a population. Probing the biological drivers for these heterogeneities, often as a function of time, forms the basis of many of the current uses of network analysis in the behavioural sciences. This special issue on social networks brings together a diverse group of practitioners whose study systems range from social insects over reptiles to birds, cetaceans, ungulates and primates in order to illustrate the wide-ranging applications of network analysis.

Abstract: A central question in the study of predator–prey relationships is to what extent prey behaviour is determined by avoidance of predators. Here, we test whether the long-term risk of encountering lions and the presence of lions in the vicinity influence the behaviour of large African herbivores at waterholes through avoidance of high-risk areas, increases in group size, changes in temporal niche or changes in the time spent in waterhole areas. In Hwange National Park, Zimbabwe, we monitored waterholes to study the behaviour of nine herbivore species under different risks of encountering lions. We radio-collared 26 lions in the study area which provided the opportunity to monitor whether lions were present during observation sessions and to map longer-term seasonal landscapes of risk of encountering lions. Our results show that the preferred prey species for lions (buffalo, kudu and giraffe) avoided risky waterholes. Group size increased as encounter risk increased for only two species (wildebeest and zebra), but this effect was not strong. Interestingly, buffalo avoided the hours of the day which are dangerous when the long-term and short-term risks of encountering lions were high, and all species showed avoidance of waterhole use at night times when lions were in the vicinity. This illustrates well how prey can make temporal adjustments to avoid dangerous periods coinciding with predator hunting. Additionally, many herbivores spent more time accessing water to drink when the long- and short-term risks of encountering lions were high, and they showed longer potential drinking time when the long-term risk of encountering lions was high, suggesting higher levels of vigilance. This study illustrates the diversity of behavioural adjustments to the risk of encountering a predator and how prey respond differently to temporal variations in this risk.

Abstract: We use two novel techniques to analyze association patterns in a group of wild spider monkeys (Ateles geoffroyi) studied continuously for 8 years. Permutation tests identified association rates higher or lower than chance expectation, indicating active processes of companionship and avoidance as opposed to passive aggregation. Network graphs represented individual adults as nodes and their association rates as weighted edges. Strength and eigenvector centrality (a measure of how strongly linked an individual is to other strongly linked individuals) were used to quantify the particular role of individuals in determining the network's structure. Female–female dyads showed higher association rates than any other type of dyad, but permutation tests revealed that these associations cannot be distinguished from random aggregation. Females formed tightly linked clusters that were stable over time, with the exception of immigrant females who showed little association with any adult in the group. Eigenvector centrality was higher for females than for males. Adult males were associated mostly among them, and although their strength of association with others was lower than that of females, their association rates revealed a process of active companionship. Female–male bonds were weaker than those between same-sex pairs, with the exception of those involving young male adults, who by virtue of their strong connections both with female and male adults, appear as temporary brokers between the female and male clusters of the network. This analytical framework can serve to develop a more complete explanation of social structure in species with high levels of fission–fusion dynamics.

Abstract: Flight initiation distance (FID), the distance at which an organism begins to flee an approaching threat, is an important component of antipredator behavior and a potential indicator of an animal’s perception of threat. In a field study on parrotfishes, we tested the predictions that FID in response to a diver will increase with body size, a correlate of reproductive value, and with experience of threat from humans. We studied a broad size range in four species on fringing reefs inside and outside the Barbados Marine Reserve. We used the Akaike's Information Criterion modified for small sample sizes (AICc) and model averaging to select and assess alternative models. Body size, reserve protection, and distance to a refuge, but not species, had strong support in explaining FID. FID increased with body size and generally remained two to ten times fish total length. FID was greater outside the reserve, especially in larger fish. Although we were not able to completely rule out other effects of size or reserve, this study supports predictions of an increase in FID with reproductive value and threat from humans.

Abstract: Ant colonies are factories within fortresses (Oster and Wilson 1978). They run on resources foraged from an outside world fraught with danger. On what basis do individual ants decide to leave the safety of the nest? We investigated the relative roles of social information (returning nestmates), individual experience and physiology (lipid stores/corpulence) in predicting which ants leave the nest and when. We monitored Temnothorax albipennis workers individually using passive radio-frequency identification technology, a novel procedure as applied to ants. This method allowed the matching of individual corpulence measurements to activity patterns of large numbers of individuals over several days. Social information and physiology are both good predictors of when an ant leaves the nest. Positive feedback from social information causes bouts of activity at the colony level. When certain social information is removed from the system by preventing ants returning, physiology best predicts which ants leave the nest and when. Individual experience is strongly related to physiology. A small number of lean individuals are responsible for most external trips. An individual’s nutrient status could be a useful cue in division of labour, especially when public information from other ants is unavailable.

Abstract: Several recent studies have tested the hypothesis that song quality in adult birds may reflect early developmental conditions, specifically nutritional stress during the nestling period. Whilst all of these earlier studies found apparent links between early nutritional stress and song quality, their results disagree as to which aspects of song learning or production were affected. In this study, we attempted to reconcile these apparently inconsistent results. Our study also provides the first assessment of song amplitude in relation to early developmental stress and as a potential cue to male quality. We used an experimental manipulation in which the seeds on which the birds were reared were mixed with husks, making them more difficult for the parents to obtain. Compared with controls, such chicks were lighter at fledging; they were thereafter placed on a normal diet and had caught up by 100 days. We show that nutritional stress during the first 30 days of life reduced the birds’ accuracy of song syntax learning, resulting in poorer copies of tutor songs. Our experimental manipulations did not lead to significant changes in song amplitude, song duration or repertoire size. Thus, individual differences observed in song performance features probably reflect differences in current condition or motivation rather than past condition.

Abstract: The adjustment to deal with intragroup food competition is probably the most plausible explanation of high levels of fission–fusion dynamics. However, studies did not always support expected relations between food availability, ranging costs, and subgroup size. We used several levels of analysis differing in the time and spatial scale in order to investigate this explanation in spider monkeys. In our study, subgroups were larger when food availability was higher across most levels of analyses used. We also found a fine-scale adjustment: compared to the food patch previously visited, spider monkeys traveled to larger patches just after fusions. This was not without an immediate travel cost: the interpatch distance and travel time after a fusion were longer than that before the fusion. This rapid adjustment shows the flexibility that fission–fusion dynamics can offer. Spider monkeys are in large subgroups only when food conditions are favorable, as evidenced by the fact that at all the other time-scale levels larger subgroups did not experience greater ranging costs than smaller subgroups. Our results indicate that on the whole spider monkeys successfully minimize ranging costs by fission and fusion of subgroups.

Abstract: Studies of innate colour preference and learning ability have focused on differences at the species level, rather than variation among populations of a single species. Initial strength and persistence of colour preferences are likely to affect colour choices of naive flower visitors. We therefore study the influence of both the strength and persistence of innate colour preference (for blue) on an operant learning task (associating food reward with yellow flowers) in two populations of the bumblebee Bombus terrestris. We found that both strength and persistence of blue preference differed significantly between populations: B. terrestris dalmatinus had a weaker and less persistent blue preference than B. terrestris audax. These differences in preference also influenced learning performance. Considering only landing behaviours, one-trial learning occurred in the majority (73%) of bees, and was achieved sooner in B. terrestris dalmatinus because of its weaker blue preference. However, compared to landing behaviours the relative frequency of approach flights to rewarding and unrewarding flower types changed more slowly with task experience in both populations. When considering both approaches and landings, the rate of learning, following the first rewarded learning trial, was faster in B. terrestris audax than B. terrestris dalmatinus. However, the net effects of population differences in blue preference and learning dynamics result in similar final levels of task performance. Our results provide new evidence of behavioural differences among isolated populations within a single species, and raise intriguing questions about the ecological significance and adaptive nature of colour preference.

Abstract: The antipredator behaviour of prey organisms is shaped by a series of threat-sensitive trade-offs between the benefits associated with successful predator avoidance and a suite of other fitness-related behaviours such as foraging, mating and territorial defence. Recent research has shown that the overall intensity of antipredator response and the pattern of threat-sensitive trade-offs are influenced by current conditions, including variability in predation risk over a period of days to weeks. In this study, we tested the hypothesis that long-term predation pressure will likewise have shaped the nature of the threat-sensitive antipredator behaviour of wild-caught Trinidadian guppies (Poecilia reticulata). Female guppies were collected from two populations that have evolved under high- and low-predation pressure, respectively, in the Aripo River, Northern Mountain Range, Trinidad. Under laboratory conditions, we exposed shoals of three guppies to varying concentrations of conspecific damage-released chemical alarm cues. Lower Aripo (high-predation) guppies exhibited the strongest antipredator response when exposed to the highest alarm cue concentration and a graded decline in response intensity with decreasing concentrations of alarm cue. Upper Aripo (low-predation) guppies, however, exhibited a nongraded (hypersensitive) response pattern. Our results suggest that long-term predation pressure shapes not only the overall intensity of antipredator responses of Trinidadian guppies but also their threat-sensitive behavioural response patterns.

Abstract: Polymorphism frequently correlates with specialized labor in social insects, but extreme morphologies may compromise behavioral flexibility and thus limit caste evolution. The ant genus Pheidole has dimorphic worker subcastes in which major workers appear limited due to their morphology to performing defensive or trophic functions, thus providing an ideal model to investigate specialization and plasticity. We examined worker morphology, brood-care flexibility, and subcaste ratio in 17 species of tropical twig-nesting Pheidole by quantifying nursing by major workers in natural colonies and in subcolonies lacking minors, in which we also measured brood survival and growth. Across species, majors performed significantly less brood care than minors in intact colonies, but increased rates of brood care 20-fold in subcolonies lacking minors. Brood nursed by majors had lower survival than brood tended by minors, although rates of brood growth did not vary between subcastes. Significant interspecific variation in rates of brood care by major workers did not lead to significant differences in brood growth or survival. Additionally, we did not find a significant association between the degree of major worker morphometric specialization and rates of nursing, growth, or survival of brood among species. Therefore, major workers showed reduced efficacy of brood care, but the degree of morphological specialization among species did not directly compromise task plasticity. The compact nests and all-or-nothing consequences of predation or disturbance on colony fitness may have influenced the evolution of major worker brood-care competency in twig-nesting Pheidole.

Abstract: Reproductive division of labor in social insects is accompanied by the reliable communication of individual fertility status. A central question is whether there exists a general mechanism underlying this communication system across species. The best way to produce reliable information is through physiological markers tightly associated with reproductive status. Cuticular hydrocarbons exhibit this link to individual fertility in several species of ants, bees, and wasps, and we present the first evidence for such a system in a non-Hymenopteran eusocial species. In the termite Zootermopsis nevadensis, we identified four polyunsaturated alkenes, which only occur in significant amounts on reproductives that are actively producing gametes. These compounds are either absent or only occur in small amounts in soldiers, worker-like larvae, and secondary reproductives with inactive gonads. In contrast to Hymenopteran social insects, both sexes express the reproductive peaks. The reproductive-specific hydrocarbons may promote tending behavior by worker-like larvae or act as a primer pheromone, inhibiting the reproductive development of immature conspecifics.

Abstract: The regulation of protein collection through pollen foraging plays an important role in pollination and in the life of bee colonies that adjust their foraging to natural variation in pollen protein quality and temporal availability. Bumble bees occupy a wide range of habitats from the Nearctic to the Tropics in which they play an important role as pollinators. However, little is known about how a bumble bee colony regulates pollen collection. We manipulated protein quality and colony pollen stores in lab-reared colonies of the native North American bumble bee, Bombus impatiens. We debut evidence that bumble bee colony foraging levels and pollen storage behavior are tuned to the protein quality (range tested: 17–30% protein by dry mass) of pollen collected by foragers and to the amount of stored pollen inside the colony. Pollen foraging levels (number of bees exiting the nest) significantly increased by 55%, and the frequency with which foragers stored pollen in pots significantly increased by 233% for pollen with higher compared to lower protein quality. The number of foragers exiting the nest significantly decreased (by 28%) when we added one pollen load equivalent each 5 min to already high intranidal pollen stores. In addition, pollen odor pumped into the nest is sufficient to increase the number of exiting foragers by 27%. Foragers directly inspected pollen pots at a constant rate over 24 h, presumably to assess pollen levels. Thus, pollen stores can act as an information center regulating colony-level foraging according to pollen protein quality and colony need.

Abstract: In social insects, the decision to exploit a food source is made both at the individual (e.g., a worker collecting a food item) and colony level (e.g., several workers communicating the existence of a food patch). In group recruitment, the recruiter lays a temporary chemical trail while returning from the food source to the nest and returns to the food guiding a small group of nestmates. We studied how food characteristics influence the decision-making process of workers changing from individual retrieving to group recruitment in the gypsy ant Aphaenogaster senilis. We offered field colonies three types of prey: crickets (cooperatively transportable), shrimps (non-transportable), and different quantities of sesame seeds (individually transportable). Colonies used group recruitment to collect crickets and shrimps, as well as seeds when they were available in large piles, while small seed piles rarely led to recruitment. Foragers were able to “measure” food characteristics (quality, quantity, transportability), deciding whether or not to recruit, accordingly. Social integration of individual information about food emerged as a colony decision to initiate or to continue recruitment when the food patch was rich. In addition, group recruitment allowed a fast colony response over a wide thermal range (up to 45°C ground temperature). Therefore, by combining both advantages of social foraging (group recruitment) and thermal tolerance, A. senilis accurately exploited different types of food sources which procured an advantage against mass-recruiting and behaviorally dominant species such as Tapinoma nigerrimum and Lasius niger.

Abstract: Despite the importance of acquired predator recognition in mediating predator–prey interactions, we know little about the specific characteristics that prey use to distinguish predators from non-predators. Recent experiments with mammals and fish indicate that some prey lacking innate predator recognition have the ability to display anti-predator responses upon their first encounter with those predators if they are similar to predators that the prey has recently learned to recognize. This phenomenon is referred to as generalization of predator recognition. In this experiment, we documented for the first time that larval amphibians (woodfrog, Rana sylvatica) have the ability to generalize the recognition of known predators to closely related novel predators. Moreover, we demonstrated that this ability is dependent on the level of risk associated with the known predator. When red-bellied newt, Cynops pyrrhogaster (known predator), was paired with simulated low risk, tadpoles displayed fright responses to newts and novel tiger salamanders, Ambystoma tigrinum, but not to novel African clawed frogs, Xenopus laevis. However, when the newt was paired with simulated high risk, tadpoles generalized their responses to both tiger salamanders and African clawed frogs. Larval anurans seem to have a wider generalization frame than other animals.

Abstract: We determined annual male reproductive success in black bears (Ursus americanus) using DNA and field data from two populations in New Mexico. We identified the likely father for 78 of 120 genotyped cubs born during 1994–2000 and calculated reproductive success for 102 males known or presumed present within designated mating regions. Age was a strong determinant of reproductive success. The observed peak in reproductive success occurred at roughly the same intermediate age (10 years) as milder peaks in body size characteristics (11–12 years) and frequency of bear-inflicted wounds (13 years), suggesting body size and fighting might be important for mating. Success was negatively associated with the distance between home range centers of males and mothers. Success of young males (<7 years old) was also negatively associated with mature male (≥7 years old) density, and increasing density shifted the peak age of reproduction higher. The dispersed distribution of females likely limited the capacity of large mature males to dominate reproduction; therefore, success was determined by a complex set of variables.

Abstract: Authors of socioecological models propose that food distribution affects female social relationships in that clumped food resources, such as fruit, result in strong dominance hierarchies and favor coalition formation with female relatives. A number of Old World monkey species have been used to test predictions of the socioecological models. However, arboreal forest-living Old World monkeys have been understudied in this regard, and it is legitimate to ask whether predominantly arboreal primates living in tropical forests exhibit similar or different patterns of behavior. Therefore, the goal of our study was to investigate female dominance relationships in relation to food in gray-cheeked mangabeys (Lophocebus albigena). Since gray-cheeked mangabeys are largely frugivorous, we predicted that females would have linear dominance hierarchies and form coalitions. In addition, recent studies suggest that long food site residence time is another important factor in eliciting competitive interactions. Therefore, we also predicted that when foods had long site residence times, higher-ranking females would be able to spend longer at the resource than lower-ranking females. Analyses showed that coalitions were rare relative to some other Old World primate species, but females had linear dominance hierarchies. We found that, contrary to expectation, fruit was not associated with more agonism and did not involve long site residence times. However, bark, a food with a long site residence time and potentially high resource value, was associated with more agonism, and higher-ranking females were able to spend more time feeding on it than lower-ranking females. These results suggest that higher-ranking females may benefit from higher food and energy intake rates when food site residence times are long. These findings also add to accumulating evidence that food site residence time is a behavioral contributor to female dominance hierarchies in group-living species.

Abstract: We investigated the natural dynamics in a sexual signal that combines different call components and explored the role of call complexity in sexual selection using a neotropical frog. Male tungara frogs, Physalaemus pustulosus, facultatively add up to seven short, multi-harmonic components (chucks) to the simple form of their calls (whines). Female tungara frogs are preferentially attracted to whines with chucks over whines without chucks, and males also call more in response to calls containing chucks. Because acoustic predators prefer complex calls, in the context of simple (no chucks) versus complex (any number of chucks) calls, the variably complex call appears to have evolved in response to the opposing selective forces of natural and sexual selection. There is no evidence, however, for the function of increasing the number of chucks within complex calls. We tested two aspects of increasing call complexity: natural patterns of use of call types in males and how both sexes respond to variation in multi-chuck calls. Males incrementally change call complexity by the addition or subtraction of a single chuck and usually do not produce more than two chucks. Variation in call complexity, for calls with at least one chuck, does not influence response calling in males or phonotaxis in females. Our results suggest that one reason for not increasing call complexity beyond a single chuck is the diminishing effectiveness on the responses of both sexes.

Abstract: A common explanation for hunting in groups is that doing so yields a greater per capita caloric benefit than hunting solitarily. This is logical for social carnivores, which rely exclusively on meat for energy, but arguably not for omnivores, which obtain calories from either plant or animal matter. The common chimpanzee, Pan troglodytes, is one of the few true omnivores that regularly hunts in groups. Studies to date have yielded conflicting data regarding the payoffs of group hunting in chimpanzees. Here, we interpret chimpanzee hunting patterns using a new approach. In contrast to the classical assumption that hunting with others maximizes per capita caloric intake, we propose that group hunting is favored because it maximizes an individual’s likelihood of obtaining important micronutrients that may be found in small quantities of meat. We describe a mathematical model demonstrating that group hunting may evolve when individuals can obtain micronutrients more frequently by hunting in groups than by hunting solitarily, provided that group size is below a certain threshold. Twenty five years of data from Gombe National Park, Tanzania are consistent with this prediction. We propose that our ‘meat-scrap’ hypothesis is a unifying approach that may explain group hunting by chimpanzees and other social omnivores.

Abstract: Offspring survival can be influenced by resources allocated to eggs, which in turn may be affected by the environmental factors the mother experiences during egg formation. In this study, we investigated whether experimentally elevated social interactions and number of neighbouring pairs influence yolk composition of collared flycatchers (Ficedula albicollis). Social challenge was simulated by presentation of a conspecific female. Experimental females spent more time near the cage and produced eggs with higher androgen concentration, but local breeding density did not affect yolk androgen level. Moreover, we found that females exposed to more intra-specific interactions and those that bred at higher density produced eggs with smaller yolk. These females may be more constrained in foraging time due to more frequent social encounters, and there might be increased competition for food at areas of higher density. In contrast, the present study did not reveal any evidence for the effect of social environment on yolk antioxidant and immunoglobulin levels. However, we found that yolk lutein and immunoglobulin concentrations were related to the female’s H/L ratio. Also, yolk lutein and α-tocopherol levels showed a seasonal increase and were positively related to the female’s plasma carotenoid level. Mothers may incur significant costs by transferring these compounds into the eggs, thus only females in good physiological condition and those that lay eggs later, when food is probably more abundant, could allocate higher amounts to the eggs without compromising their defence mechanisms. Our results suggest that environmental circumstances during egg formation can influence conditions for embryonic development.

Abstract: Female seed beetles, Callosobruchus maculatus, mate multiply even though association with males and copulations carry costs, such as injury to the genital tract. Multiple mating (polyandry) may, however, offset these costs through the acquisition of food and water, two material benefits hypothesized to be obtained from the large ejaculates produced by males. The material benefits hypothesis can be tested by increasing female access to nutrients and water, with the prediction that female mating frequency will decrease as copulation is no longer required to derive these materials. Females were given water, 5% sugar-water or baker's yeast, and were compared with females deprived of these. We presented females with virgin males daily for 8 days and recorded female mating frequency, survivorship, and fecundity. Females provided with water and sugar-water decreased mating frequency. Thus, water, rather than nutrients in the ejaculate, appears to be important to females of this species. In addition, both life span and fecundity were extended for females in the sugar-water and water treatments. Since water is scarce in the arid environment in which this species is found, we conclude that polyandry provides material benefits to females that may offset some of the costs of associating with males.

Abstract: We studied two courtship displays of male peafowl (Pavo cristatus), focusing particularly on male orientation relative to the position of the sun. During the “wing-shaking” display, females were generally behind the displaying male, and male orientation with respect to the position of the sun was not significantly different from random. However, during the pre-copulatory “train-rattling” display, males were on average directed at about 45° to the right of the sun azimuth with the female positioned directly in front, suggesting that this behaviour is involved in the communication of a visual signal. A model presentation experiment confirmed that courting peacocks were more likely to perform the train-rattling display when the female was on the sunny side of their erect train, but more likely to perform wing-shaking behaviour when the female was on the shaded side of the male. This study underscores the importance of visual signalling in peafowl courtship, and we suggest that an angle of about 45° relative to the sun may allow males to enhance the appearance of their iridescent eyespot feathers.

Abstract: In societies characterized by a high degree of fission-fusion dynamics, individuals adjust their grouping patterns according to the shifting balance of costs and benefits associated with grouping. This study examines influences on fission-fusion dynamics for dusky dolphins (Lagenorhynchus obscurus) in Admiralty Bay, New Zealand. This area is an important foraging habitat for dusky dolphins during the winter and spring. Admiralty Bay has little predation risk, but nearshore mussel farms may infringe on available habitat. I used generalized estimating equations to determine the influences of coordinated foraging, predation risk, and presence of mussel farms on party size, rate of fission-fusion, and behavioral state. I conducted 168 boat-based group focal follows totaling 168 h. The proportion of individuals observed foraging was positively related to party size and rate of party fusion. Resting had no effect on party size and did not vary according to location. Near mussel farms, traveling decreased, and rate of party fission decreased. I conclude that (1) coordinated foraging strategies are a primary influence on fission-fusion dynamics within this population, (2) dolphins may respond to decreased predation risk by not adjusting party size or location during resting, and (3) areas near mussel farms are not used for traveling.

Abstract: Animals’ contest performance is influenced by their recent contest experiences. This influence could either be exerted by individuals re-estimating their own fighting ability (self-assessment) or by their opponents responding to status-related cues (social-cue mechanism) or both. Individuals of Kryptolebias marmoratus, a hermaphroditic killifish, were given different contest experiences to examine how two opponents’ prior experiences combined to determine their contest interaction and to test both of these mechanisms as potential causes of the observed experience effect. Our data showed that losers’ decisions to retreat at different stages of a contest were influenced by their own but not by the winners’ contest experience—a result consistent with self-assessment but not with the social-cue mechanism. An association between the fish initiating and winning contests thus probably arose because both were correlated with an individual’s assessment of its fighting ability, but not because initiating contests made opponents more inclined to retreat.

Abstract: Adult male chacma baboons (Papio hamadryas ursinus) form preferential associations, or friendships, with particular lactating females. Males exhibit high levels of affiliative contact with their friends’ infants and defend them from potentially infanticidal attacks (Palombit et al. 1997). Little is known about males’ associations with juveniles once they have passed the period of infanticidal risk. We conducted an observational, experimental, and genetic study of adult male and juvenile chacma baboons in the Moremi Reserve, Botswana. We identified preferential associations between males and juveniles and used behavioral data and a playback experiment to explore whether those associations have potential fitness benefits for juveniles. We determined whether males preferentially invest in care of their own offspring. We also determined how often males invest in care of their former friends’ offspring. The majority of juveniles exhibited preferential associations with one or two males, who had almost always been their mother’s friend during infancy. However, in only a subset of these relationships was the male the actual father, in part because many fathers died or disappeared before their offspring were weaned. Male caretakers intervened on behalf of their juvenile associates in social conflicts more often than they intervened on behalf of unconnected juveniles, and they did not appear to differentiate between genetic offspring and unrelated associates. Playbacks of juveniles’ distress calls elicited a stronger response from their caretakers than from control males. Chacma males may provide care to unrelated offspring of former friends because the costs associated with such care are low compared with the potentially high fitness costs of refusing aid to a juvenile who is a possible offspring.

Abstract: The environment is profoundly important in shaping many aspects of animal phenotype, including courtship and mating behaviours. Courtship displays rely upon the transmission of visual information from the signaller to the receiver, which means they are likely to be less effective in visually poor conditions such as at low light or in turbid ecosystems. One might therefore predict that in visually poor environments it would be beneficial for individuals to plastically adjust their mating behaviour to maximise mating success. Here, we investigate the impact of the developmental and current visual environment (light intensity) upon male mating behaviour in the Trinidadian guppy Poecilia reticulata. Male guppies have two different mating tactics: They can court females with a visual sigmoid display or attempt to circumvent female choice by attempting a non-consensual copulation (gonapodium thrust). We reared juvenile guppies in low light and relatively high light intensities for 5 months before observing individual males for mating behaviour in both light conditions. We found that the current light environment is important in determining the frequency of both sigmoidal courtship displays and non-consensual copulation attempts. Males increase the frequency of sigmoidal displays at relatively high light and increase non-consensual mating attempts at low light, suggesting that males compensate for poor visual conditions via an adjustment in tactics. We also find a significant correlation in courtship effort between the different light environments, suggesting that there is individual consistency across time and context for this trait. Developmental environment was less important. However, we found that fish reared at lower light intensities continued to employ sigmoidal displays despite the poor current visual environment. Our data show that male mating behaviour is phenotypically plastic in response to recent light environment. This may have implications for understanding how animals cope with anthropogenic environmental change.

Abstract: Nitric oxide (NO) is a multifunctional signalling molecule, acting as a vasodilator, neurotransmitter, and modulator of inflammatory processes. It also participates in killing parasites, virus-infected cells, and tumor cells by formation of peroxynitrite, one of the most important initiators of the free radical damage. Uncontrolled production of NO can lead to nitrosative stress, causing damages to proteins and DNA and cell injury and death. Determination of NO production in animals is potentially informative for description of individual variation in physiological condition, health state, and work load; however, it’s potential in ecophysiological research has remained almost totally unexplored. Here, we describe application of a simple, precise, and inexpensive spectrophotometric assay for determination of NO production from 5 to 10 μL plasma samples of passerine birds. The method is based on estimation of concentrations of nitrate and nitrite—the stable end products of nitric oxide oxidation. The principle of the assay is reduction of nitrate to nitrite by copper-coated cadmium granules, followed by color development with Griess reagent. NO production in captive greenfinches (Carduelis chloris L.) was significantly repeatable over 6-day period (r = 0.35). Injection of an inflammatory agent phytohemagglutinin into wing web resulted in 21% higher levels of NO production at the third day after treatment as compared to saline-injected birds. These findings, consistently with further evidence from veterinary and biomedical literature, suggest that measuring NO production appears an efficient and robust tool for monitoring individual condition and assessment of the magnitude of innate immune response, pathogenicity of infections, and physical effort. We propose that this assay, which is easily applicable in field studies, has an excellent potential in ecophysiological research, particularly in the rapidly developing fields of immunoecology and conservation physiology.

Abstract: Social nesting behaviour is commonly associated with high prevalence and intensity of parasites in intraspecific comparisons. Little is known about the effects of interspecific host breeding density for parasite intensity in generalist host–parasite systems. Darwin’s small tree finch (Camarhynchus parvulus) on Santa Cruz Island, Galapagos Islands, nests in both heterospecific aggregations and at solitary sites. All Darwin finch species on Santa Cruz Island are infested with larvae of the invasive blood-sucking fly Philornis downsi. In this study, we test the prediction that total P. downsi intensity (the number of parasites per nest) is higher for nests in heterospecific aggregations than at solitary nests. We also examine variation in P. downsi intensity in relation to three predictor variables: (1) nest size, (2) nest bottom thickness and (3) host adult body mass, both within and across finch species. The results show that (1) total P. downsi intensity was significantly higher for small tree finch nests with many close neighbours; (2) finches with increased adult body mass built larger nests (inter- and intraspecific comparison); (3) parasite intensity increased significantly with nest size across species and in the small tree finch alone; and (4) nest bottom thickness did not vary with nest size or parasite intensity. These results provide evidence for an interaction between social nesting behaviour, nest characteristics and host mass that influences the distribution and potential impact of mobile ectoparasites in birds.

Abstract: The regulation of protein collection through pollen foraging plays an important role in pollination and in the life of bee colonies that adjust their foraging to natural variation in pollen protein quality and temporal availability. Bumble bees occupy a wide range of habitats from the Nearctic to the Tropics in which they play an important role as pollinators. However, little is known about how a bumble bee colony regulates pollen collection. We manipulated protein quality and colony pollen stores in lab-reared colonies of the native North American bumble bee, Bombus impatiens. We debut evidence that bumble bee colony foraging levels and pollen storage behavior are tuned to the protein quality (range tested: 17–30% protein by dry mass) of pollen collected by foragers and to the amount of stored pollen inside the colony. Pollen foraging levels (number of bees exiting the nest) significantly increased by 55%, and the frequency with which foragers stored pollen in pots significantly increased by 233% for pollen with higher compared to lower protein quality. The number of foragers exiting the nest significantly decreased (by 28%) when we added one pollen load equivalent each 5 min to already high intranidal pollen stores. In addition, pollen odor pumped into the nest is sufficient to increase the number of exiting foragers by 27%. Foragers directly inspected pollen pots at a constant rate over 24 h, presumably to assess pollen levels. Thus, pollen stores can act as an information center regulating colony-level foraging according to pollen protein quality and colony need.

Abstract: The availability of food resources changes over time and space, and foraging animals are constantly faced with choices about how to respond when a resource becomes depleted. We hypothesise that flying insects like bees discover new food sources using an optimal Levy flight searching strategy and odour-mediated anemotaxis, as well as visual cues. To study these searching patterns, foraging honeybees were trained to a scented feeder which was then removed. Two new unrewarding feeders, or ‘targets’, were then positioned up- and downwind of the original location of the training feeder. The subsequent flight patterns of the bees were recorded over several hundred metres using harmonic radar. We show that the flight patterns constitute an optimal Levy flight searching strategy for the location of the training feeder, a strategy that is also optimal for the location of alternative food sources when patchily distributed. Scented targets that were positioned upwind of the original training feeder were investigated most with the numbers of investigations declining with increasing distance from the original feeder. Scented targets in downwind locations were rarely investigated and unscented targets were largely ignored, despite having the same visual appearance as the rewarding training feeder.