The honey bee shaking signal: function and design of a modulatory communication signal

This study explores the meaning and functional design of a modulatory communication signal, the honey bee shaking signal, by addressing five questions: (I) who shakes, (II) when do they shake, (III) where do they shake, (IV) how do receivers respond to shaking, and (V) what conditions trigger shaking. Several results confirm the work of Schneider (1987) and Schneider et al. (1986a): (I) most shakers were foragers (at least 83%); (II) shaking exhibited a consistent temporal pattern with bees producing the most signals in the morning (0810–1150 hours) just prior to a peak in waggle dancing activity; and (IV) bees moved faster (by 75%) after receiving a shaking signal. However, this study differs from previous work by providing a long-term, temporal, spatial, and vector analysis of individual shaker behavior. (III) Bees producing shaking signals walked and delivered signals in all areas of the hive, but produced the most shaking signals directly above the waggle dance floor. (IV) Bees responded to the signal by changing their direction of movement. Prior to receiving a signal, bees selected from the waggle dance floor moved, on average, towards the hive exit. After receiving a signal, some bees continued moving towards the exit but others moved directly away from the exit. During equivalent observation periods, non-shaken bees exhibited a strong tendency to move towards the hive exit. (V) Renewed foraging activity after food dearth triggered shaking signals, and, the level of shaking is positively correlated with the duration of food dearth. However, shaking signal levels also increased in the morning before foraging had begun and in the late afternoon after foraging had ceased. This spontaneous afternoon peak has not previously been reported. The shaking signal consequently appears to convey the general message “reallocate labor to different activities” with receiver context specifying a more precise meaning. In the context of foraging, the shaking signal appears to activate (and perhaps deactivate) colony foraging preparations. The generally weak response elicited by modulatory signals such as the shaking signal may result from a high receiver response threshold which allows the receiver to integrate multiple sources of information and which thereby increases the probability that receiver actions will be appropriate to colony needs. Received: 21 March 1997 / Accepted after revision: 30 August 1997     

Bird song as a signal of aggressive intent

A central question in animal communication research concerns the reliability of animal signals. The question is particularly relevant to aggressive communication, where there often may be advantages to signaling an exaggerated likelihood of attack. We tested whether aggressive signals are indeed reliable signals of attack in song sparrows (Melospiza melodia). We elicited aggressive signaling using a 1-min playback on a male’s territory, recorded the behavior of the male for 5 min, and then gave him the opportunity to attack a taxidermic mount of a song sparrow associated with further playback. Twenty subjects attacked the mount and 75 did not. Distance to the speaker was a significant predictor of attack for both the initial recording period and the 1 min before attack. For the initial recording period, none of the measures of singing behavior that we made was a significant predictor of attack, including song-type matching, type-switching frequency, and song rate. For the 1-min period immediately before attack, only the number of low amplitude “soft songs” was a significant predictor of attack. Although most aggressive signals contained little information on attack likelihood, as some models suggest should be the case, the unreliability of these signals was not caused by convergence of individuals on a single signaling strategy, as those models argue should occur.     

The presence of females modulates the expression of a carotenoid-based sexual signal

Several environmental factors have been shown to shape the pattern of investment of carotenoids into the expression of sexual signals. Surprisingly, the impact of the social environment has been neglected. If a carotenoid-based sexual trait is used by females to choose a mate or by males to assess the quality of potential competitors for mates, males, in the presence of females, should upregulate expression of the trait. We tested this hypothesis in male zebra finches that were housed either with females or in a male-only social environment. Additionally, to investigate whether the social modulation of the expression of a sexual trait depends on the physiological need of carotenoids, we challenged half of the males with Escherichia coli lipopolysaccharide (LPS). We found that the social environment modulated the expression of bill color, with males kept in the presence of females harboring a redder bill at the end of the 3 weeks that the experiment lasted. Males injected with LPS showed duller bill color regardless of the presence of females, and social interactions with females result in upregulated bill color similarly for phosphate-buffered saline (PBS) and LPS males. Thus, social environment and immune activation had an additive effect on the expression of bill color. The effect of social environment on plasma carotenoids was less clear. Indeed, a first replicate of the entire experiment showed that both immune challenge and social context affected bill color, with a negative effect of immune challenge and a positive effect of the presence of females on circulating carotenoids. However, a second replicate of the experiment showed only a negative effect of the immune challenge. These results, therefore, suggest that the social environment can affect the expression of carotenoid-based sexual traits under both benign and carotenoid-demanding conditions. Whatever the signaling function of bill color (female mate choice or male–male competition for mates), the observed flexibility may be adaptive because the expression of the signal can be modulated depending on the expected rewards or costs associated with the presence or absence of females. Nevertheless, the mechanisms underlying such an effect are still unknown.     

To whom it may concern: the transmission and function of chemical signals in Lemur catta

The goals of this study were to investigate the transmission and possible functions of chemical signals in intragroup communication among ring-tailed lemurs, Lemur catta. In particular, I examined the effects of sex on these processes because sexual selection theory predicts specific functions for chemical signals. I recorded all interactions with 214 scent marks of 11 male and 9 female ring-tailed lemurs during the first 10 min following their deposition. I found that 62% of these scent marks were investigated with a median latency of 30 s and that 89% of investigated scents were also counter-marked by the receiver. The type of focal scent mark (male or female anogenital and male antebrachial mark) had a significant effect on both the timing and type of response. Males investigated and counter-marked female scents more often than vice versa, but significant second-order transitions suggested that the behavior of an animal was not only influenced by the immediately preceding scent mark and that a scent is not completely masked by a counter-mark. There was no evidence for an audience effect, and only social rank of female senders had an effect on receivers. Variation in the response of receivers across reproductive seasons as a function of senders' sex indicated that female scents may function in mate attraction and competition among females, whereas male scents may be primarily used in intrasexual competition. Three main conclusions emerged. First, the exchange of olfactory signals within groups was highly structured and surprisingly efficient. Second, olfactory signals may constitute general mesasages whose transfer is partly controlled by the receiver. Finally, sexual selection theory provides a useful theoretical framework for functional examinations of mammalian olfactory communication. Received: 13 June 1997 / Accepted after revision: 20 March 1998     

Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.     

Risk taking during parental care: a test of three hypotheses applied to the pied flycatcher

According to life-history theory, there will often be a conflict between investment in current versus future reproduction. If a predator appears during breeding, parents must make a compromise between ensuring the growth and survival of offspring (nest defence, feeding and brooding of young), and reducing the risk of predation to ensure their own survival. We model three hypotheses for the outcome of this conflict which are particularly relevant for altricial birds. They are not mutually exclusive, but focus on different costs and benefits. (1) Parental investment is determined by the parents’ own risk of predation. This hypothesis predicts that a lone parent should take smaller risks than a parent that has a mate. (2) Parental investment is related to the reproductive value of the offspring: Parents are predicted to take greater risks for larger broods, larger-sized or older offspring. (3) Finally, we present the new hypothesis that parental investment is related to the harm that offspring would suffer during a period of no parental care (incubation, brooding, feeding). This hypothesis predicts that parents should take greater risks for younger offspring, or for offspring in poorer condition, because the marginal benefit of parental care is largest in such cases. Hence, one may also expect that lone parents should take greater risks than two parents because their offspring are more in need of care. We tested these hypotheses on the pied flycatcher (Ficedula hypoleuca) by presenting a stuffed predator of the parents (a sparrowhawk, Accipiter nisus) close to the nest when parents were feeding the young. Risk taking was measured as the time that elapsed until the first visit to the nest. Most support was found for the ‘‘harm to offspring’’ hypothesis. Previous studies have usually measured the intensity of nest defence against typical nest predators, and have found evidence for the ‘‘reproductive value of offspring’’ hypothesis. However, our model predicts that the importance of the reproductive value of the offspring should decrease relative to the harm that offspring would suffer if they were not cared for when the predator type changes from a nest predator to a predator of adults, and when conditions for breeding turn from good to bad. Received: 13 April 1995/Accepted after revision: 11 March 1996     

Female territoriality and the function of scent-marking in a monogamous antelope (Oreotragus oreotragus)

In monogamous species, females often choose between males according to the quality of the territories they defend, but the extent to which females themselves contribute to territory defence is frequently underestimated. Here we test for differences in male and female roles during paired scent-marking bouts, a key component of territorial defence, in a monogamous antelope. In two populations (Kenya, Zimbabwe) of klipspringer, Oreotragus oreotragus, both males and females usually scent-marked at the same site, but there were significant differences between sexes in terms of investment within bouts. Females initiated most bouts, thus dictating the marking strategy of the pair. Males initiated relatively few bouts, but deposited more scent marks per bout than females and were usually the last to scent-mark before leaving the site; they marked on the same branches as the female and thus overmarked her scent. Both sexes deposited more marks during paired than solo visits. Immediately preceding and following scent-marking bouts, males approached females and females left males more often than expected. Female scent-marking rates were higher when they were receptive than at other times, and this increase was matched by elevated marking rates of males. Females may increase marking rates when they are receptive in order to test the quality of their mate or to incite male competition. However, these ideas are unlikely to explain female scent-marking behaviour outside the mating season, which appears to be related primarily to territorial defence. We suggest that these differences in investment in scent-marking bouts are consistent with predictions that females may be autonomously territorial and that overmarking of female scent by males is a form of mate-guarding. Received: 17 November 1999 / Received in revised form: 24 February 2000 / Accepted: 13 March 2000     

镉和铁及其交互作用对植物生长的影响

镉是植物的非必需元素，而铁是植物生长发育过程中必不可少的元素。虽然这两种元素对植物的作用是截然相反的，但是它们在植物体内存在着交互作用关系，相互影响着对方功能的发挥。供给植物足够的铁可极大地缓解植物的镉污染毒害，利于植物合成更多的叶绿素，促进光合作用进行，降低镉对氨基酸、蛋白质合成以及酶活性的影响。相反，镉污染将诱导植物的铁营养缺乏。渍水植物根表形成的铁氧化物胶膜可促进植物对镉的吸收。     

Positive association of queen number and queen-mating frequency Myrmica ants: a challenge to the genetic-variability hypotheses

Detailed knowledge of the mating system in specific social insect populations is essential for testing general evolutionary hypotheses of multiple paternity in eusocial Hymenoptera. We have studied the mating frequency of queens in a polygynous population of the red ant Myrmica sulcinodis. Genetic mother-offspring analysis showed that double mating occurred at a considerable frequency, but that the effective number of queen-mates remained close to one. After quantifying the effects of multiple maternity (polygyny) and multiple paternity (polyandry) on the genetic diversity of workers, we conclude that multiple paternity in M. sulcinodis did not evolve as an adaptation to increase genetic variation within colonies. Contrary to the predictions from `genetic variability' hypotheses, we found a positive correlation between colony-specific queen number and the average number of mates per queen. Such positive association of queen number and frequency of multiple mating was also found after analysing comparative data across six species of Myrmica ants. These results suggest that resticted dispersal of young queens may be a common factor promoting both polygyny and polyandry at the same time, and that moderate degrees of multiple mating may be an unselected consequence of (1) mating at low cost when mating occurs close to the nest and (2) mating in swarms with a highly male biased operational sex ratio. Future comparative tests of genetic-variability hypotheses should therefore not include species with such evolutionary derived mating system characteristics. Received: 30 April 1998 / Accepted after revision: 19 August 1998     

Structure and resilience of the social network in an insect colony as a function of colony size

Social interactions are critical to the organization of worker activities in insect colonies and their consequent ecological success. The structure of this interaction network is therefore crucial to our understanding of colony organization and functioning. In this paper, I study the properties of the interaction network in the colonies of the social wasp Ropalidia marginata. I find that the network is characterized by a uniform connectivity among individuals with increasing heterogeneity as colonies become larger. Important network parameters are found to be correlated with colony size and I investigate how this is reflected in the organization of work in colonies of different sizes. Finally, I test the resilience of these interaction networks by experimental removal of individuals from the colony and discuss the structural properties of the network that are related to resilience in a social network. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau, and R. James).     

PICABUE: a methodological framework for the development of indicators of sustainable development

SUMMARY

Significant interest in the concept of sustainable development exists amongst scientists, planners, policy makers and the public, and considerable effort and expenditure is made or envisaged at local, national and international levels to promote a more sustainable society. Until ‘green accounting’ and similar systems are made available and are implemented, the sustainability indicator will be the most effective tool available for monitoring progress towards a more sustainable society. Sustainability indicators are already available but are characterized by a poor or absent theoretical underpinning. This paper addresses this problem by proposing a methodological framework that can be applied to the construction of indicators of sustainable development. In order to be consistent with widely accepted definitions of sustainable development, considerations relating to the measurement of quality of life and ecological integrity are central to the methodology. The methodological framework has relevance to a variety of spatial scales and to geographically diverse areas (urban or rural, developed or developing countries) so that a suite of sustainability indicators can be produced that is tailored to the needs and resources of the indicator user, but which remains rooted firmly in the fundamental principles of sustainable development.     

The neglected scent: on the marking function of urine in Lemur catta

In ring-tailed lemurs (Lemur catta) urine deposition can be combined with two different tail configurations: (i) tail held up in an evident display (urinate tail-up, UT-up); (ii) tail slightly raised to avoid its impregnation with urine (urinate tail-down, UT-down). We used both signaller- and receiver-based approaches to search for functional differences between these two kinds of urine deposition. We predicted that UT-up might be a complex signal combining olfactory and visual cues. We carried out observations and scent tests on four captive groups of ring-tailed lemurs. Group members sniffed/licked UT-up scents more frequently than UT-down ones. Moreover, UT-up showed peak levels during the mating season whereas UT-down did not. These findings suggest that urine can play a role in intra-group reproductive communication. Lemurs more frequently performed UT-up in a few drops and UT-down in streams. Recognition experiments clearly showed that individuals can discriminate between urine of their own group and urine from a foreign group (a necessary prerequisite for the use of urine in inter-group communication). The possible function of UT-up in inter-troop communication was supported by the higher frequency of this pattern along a fence separating two of the study groups. Moreover, in the presence of a dummy, the frequency of UT-up increased significantly. In conclusion, UT-up is a complex signal with multiple characteristics. By using different sensory channels, UT-up provides different types of information (location and signaller quality) and contains multiple messages directed both at group-members and neighbouring groups....to conceal this letter, the Minister had resorted to the comprehensive and sagacious expedient of not attempting to conceal it at all (from the Purloined letter E. A. Poe, 1845)     

Intertype mark correlation function: A new tool for the analysis of species interactions

The spatial pattern of the different species in complex ecosystems reflects the underlying ecological processes. In this paper a second order moment function is proposed and tested to analyse the spatial distribution of a mark, which could be a tree characteristic such as diameter or height, between two different types of points, which could be two different tree species. The proposed function was a conditional density function based on the intertype K_rs(d) function, incorporating as test function the correlation of the marks between pairs composed of points of different types. The results obtained in simulated and real plots prove that the function is capable of revealing the scale at which spatial correlation of the mark between two types of points exists. The proposed function allows the spatial association between individuals of different species at different life stages to be identified. This analysis may reveal information on species ecology and interspecific interactions in forest ecosystems.     

Spacing and group coordination in a nocturnal primate, the golden brown mouse lemur (Microcebus ravelobensis): the role of olfactory and acoustic signals

In order to remain stable, dispersed social groups have to solve two fundamental problems: the coordination of movement and cohesiveness within a group and the spacing between the groups. Here, we investigate mechanisms involved in intra-group coordination and inter-group spacing using the golden brown mouse lemur, Microcebus ravelobensis, as a model for a nocturnal, solitary foraging mammal with a dispersed social system. By means of radiotelemetry and bioacoustics we studied the olfactory and vocal behaviour during nocturnal dispersal and reunion of five sleeping groups.All groups used 3–17 sleeping sites exclusively, suggesting a sleeping site-related territoriality and competition for them. The occurrence of olfactory and vocal behaviour showed an asymmetrical temporal distribution. Whereas marking behaviour was observed exclusively during dispersal, a particular call type, the trill, was used by all groups during reunions. Interestingly, these trills carried group-specific signatures.Our findings provide the first empirical evidence for nocturnal primates in a natural environment that olfactory signals represent an important mechanism to regulate the distribution of different groups in space, whereas acoustic signals control intra-group cohesion and coordination.     

A same-sex stepparent shortens a prebreeder's duration on the natal territory: tests of two hypotheses in Florida scrub-jays

Prebreeders of the Florida scrub-jay (Aphelocoma coerulescens) are less likely to be found on their natal territories with a same-sex stepparent than with parents or an opposite-sex stepparent. We tested two models that had been proposed to account for this sexual asymmetry. The dominance hypothesis states that stepparents perceive same-sex prebreeders as competitors, primarily for a mate, so behave aggressively toward them. The pair-formation hypothesis states that prebreeders remain home to pair eventually with the opposite-sex stepparent. Predictions from these two models were tested by analyses from a quarter-century of records on a study population at Archbold Biological Station and by new behavioral field observations at the Station. Results clearly rejected the latter and strongly supported the former hypothesis. No prediction from the pair-formation hypothesis was confirmed: no pairing by a prebreeder of either sex has ever occurred with a stepparent; remaining home was equally frequent in age-one males with stepmothers and genetic mothers, and in females with stepfathers and genetic fathers; and the same results were found in age-two prebreeders. By contrast, the data strongly supported the dominance hypothesis. Field data showed higher aggression rates by stepfathers to male prebreeders and stepmothers to female prebreeders than by parents to their same-sex genetic offspring. Fewer age-one males remained home with a stepfather than with the genetic father, and the same was found for age-one females with a stepmother and genetic mother; at age two, the effect occurred only in female prebreeders. The two hypotheses make different predictions about prebreeders with two stepparents versus with both parents, thus providing a critical test. The dominance hypothesis correctly predicted decreased duration at home by age-one males and females; data for age-two females were in the direction of predicted difference but not significant; and, as in other tests, no effect was found for age-two males. We propose that age-two male prebreeders remain home despite elevated aggression from stepfathers because these prebreeders retain the possibility of budding from or inheriting the natal territory. Received: 16 July 1997 / Accepted after revision: 26 June 1998     

Sex allocation in a monomorphic seabird with a single-egg clutch: test of the environment, mate quality, and female condition hypotheses

Sex allocation theory posits that mothers should preferentially invest in sons when environmental conditions are favorable for breeding, their mates are of high quality, or they are in good body condition. We tested these three hypotheses in rhinoceros auklets (Cerorhinca monocerata), monomorphic seabirds that lay a single-egg clutch, in 2 years that differed in environmental conditions for breeding. Results supported the environment and mate quality hypotheses, but these effects were interactive: offspring sex was independent of paternal traits in the poor year for breeding, while females mated to larger and more ornamented males reared more sons in the better year. Conversely, offspring sex was unrelated to female condition, as indexed by hatching date. We propose that good rearing conditions enable females to rear sons possessing the desirable phenotypic attributes of their mates. Results also supported two critical assumptions of sex allocation theory: (1) dimorphism in offspring condition at independence: daughters fledged with higher baseline levels of corticosterone than sons and (2) differential costs of rearing sons versus daughters: mothers rearing sons when environmental conditions were poor completed parental care in poorer condition than mothers rearing daughters in the same year and mothers rearing either sex when conditions were better. These novel results may help to explain the disparate results of previous studies of avian sex allocation.     

Exploration and exploitation of food sources by social insect colonies: a revision of the scout-recruit concept

Social insect colonies need to explore and exploit multiple food sources simultaneously and efficiently. At the individual level, this colony-level behaviour has been thought to be taken care of by two types of individual: scouts that independently search for food, and recruits that are directed by nest mates to a food source. However, recent analyses show that this strict division of labour between scouts and recruits is untenable. Therefore, a modified concept is presented here that comprises the possible behavioural states of an individual forager (novice forager, scout, recruit, employed forager, unemployed experienced forager, inspector and reactivated forager) and the transitions between them. The available empirical data are reviewed in the light of both the old and the new concept, and probabilities for the different transitions are derived for the case of the honey-bee. The modified concept distinguishes three types of foragers that may be involved in the exploration behaviour of the colony: novice bees that become scouts, unemployed experienced bees that scout, and lost recruits, i.e. bees that discover a food source other than the one to which they were directed to by their nest mates. An advantage of the modified concept is that it allows for a better comparison of studies investigating the different roles performed by social insect foragers during their individual foraging histories. Received: 29 December 1999 / Revised: 25 February 2000 / Accepted: 16 October 2000     

Food exploitation by a winter flock of greylag geese: behavioral dynamics,competition and social status

Summary To investigate the dynamics of the winter flock patch exploitation, feeding experiments were performed with 140 semi-tame, free-roaming greylag geese (Anser anser). Three different initial densities of barley were offered on a 50-m² patch: low (1600 grains/m²), intermediate (3300/m²) and high (I 1500/m²). Goose numbers on the patch, peck rates, and frequencies of agonistic encounters and of alert postures were observed by scanning the flock and in focal individuals. At low and intermediate initial food densities, peck rates decreased with food density, whereas at high food density, peck rates decreased only slightly over the feeding bout. Agonistic interactions increased as food decreased. A switch from exploitation to interference competition occurred at a threshold of approximately 900 remaining grains per square meter. With high initial food density, agonistic encounters began to increase gradually after 20–30 min. The same general patterns were observed for frequency of alertness. Family members occupied the patch for the longest time periods. The family female and the offspring fed most intensely. The family gander however, fed significantly less but was more aggressive and vigilant than all other social categories. Solitary geese spent their time on the patch doing little else but feeding and were the first to leave. It seems that high-ranking families and low-ranking singles adopt different competitive strategies, the former being superior interference competitors whereas the latter are forced scramble competitors.This study is dedicated to the memory of Konrad Lorenz Correspondence to: K. Kotrschal     

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

Behavioral games involving a clever prey avoiding a clever predator: An individual-based model of dusky dolphins and killer whales

Faced with an intermittent but potent threat, animals exhibit behavior that allows them to balance foraging needs and avoid predators and over time, these behaviors can become hard-wired adaptations with both species trying to maximize their own fitness. In systems where both predator and prey share similar sensory modalities and cognitive abilities, such as with marine mammals, the dynamic nature of predator-prey interactions is poorly understood. The costs and benefits of these anti-predator adaptations need to be evaluated and quantified based on the dynamic engagement of predator and prey. Many theoretic models have addressed the complexity of predator-prey relationships, but few have translated into testable mechanistic models. In this study, we developed a spatially-explicit, geo-referenced, individual-based model of a prototypical adult dusky dolphin off Kaikoura, New Zealand facing a more powerful, yet infrequent predator, the killer whale. We were interested in two primary objectives, (1) to capture the varying behavioral game between a clever prey and clever predator based on our current understanding of the Kaikoura system, (2) to compare evolutionary costs vs. benefits (foraging time and number of predator encounters) for an adult non-maternal dusky dolphin at various levels of killer whale-avoidance behaviors and no avoidance rules. We conducted Monte Carlo simulations to address model performance and parametric uncertainty. Mantel tests revealed an 88% correlation (426 × 426 distance matrix, km²) between observed field sightings of dusky dolphins with model generated sightings for non-maternal adult dusky dolphin groups. Simulation results indicated that dusky dolphins incur a 2.7% loss in feeding time by evolving the anti-predator behavior of moving to and from the feeding grounds. Further, each evolutionary strategy we explored resulted in dolphins incurring an additional loss of foraging time. At low killer whale densities (appearing less than once every 3 days), each evolutionary strategy simulated converged towards the evolutionary cost of foraging, that is, the loss in foraging time approached the 2.7% loss experienced by evolving near shore-offshore movement behavior. However, the highest level of killer whale presence resulted in 38% decreases in foraging time. The biological significance of these losses potentially incurred by a dusky dolphin is dependent on various factors from dolphin group foraging behavior and individual energy needs to dolphin prey availability and behavior.