Does individual personality predict male mating preference for female body size in the Trinidadian guppy?

Within populations, individual animals vary considerably in their behaviour, including mate choice and personality. There is mounting interest in the potential covariation between these two behaviours within individuals, such that personality would influence mate choice. We experimentally tested this proposition under controlled laboratory conditions using male Trinidadian guppies (Poecilia reticulata) as a model study system. We assayed repeatedly the mating preference of individual males for females based on their body size. Additionally, we assayed repeatedly two ecologically relevant personality traits in males, namely exploration of a novel environment and boldness under a simulated predation threat. Finally, we analysed whether male mating preference and personality traits were repeatable, and tested whether the personality of individual males was correlated (covaried) with their mating preference scores. Although all but one of the measures of exploration and boldness behaviour were repeatable over time, male mating preference scores were not repeatable. Measures of male exploration and boldness were not inter-correlated among individuals, suggesting the absence of a behavioural syndrome between exploration and boldness. Unexpectedly, males did not exhibit on average a significant mating preference for larger females over smaller ones; they chose randomly between the paired stimulus females. Overall, we found no compelling evidence for a relationship between individual personality traits and mating preference in male guppies, suggesting that personality does not predict mate choice, at least in our study population and under our experimental conditions. We discuss potential factors, other than male personality and body length, that might maintain inter-individual variation in male mating preferences in the guppy in the wild.     

Maladaptive behavior reinforces a recruitment bottleneck in newly settled fishes

Settlement from the plankton ends the major dispersive stage of life for many marine organisms and exposes them to intense predation pressure in juvenile habitats. This predation mortality represents a life-history bottleneck that can determine recruitment success. At the level of individual predator–prey interactions, prey survival depends upon behavior, specifically how behavior affects prey conspicuousness and evasive ability. We conducted an experiment to identify behavioral traits and performance levels that are important determinants of which individuals survive or die soon after settlement. We measured a suite of behavioral traits on late stage, pre-settlement Ward's damsel (Pomacentrus wardi) collected using light traps. These behavioral traits included two measures of routine swimming (indicators of conspicuousness) and eight measures of escape performance to a visual startle stimulus. Fish were then released onto individual patch reefs, where divers measured an additional behavioral trait (boldness). We censused each patch reef until approximately 50% of the fish were missing (~24 h), which we assumed to be a result of predation. We used classification tree analysis to discriminate survivors from fish presumed dead based on poor behavioral performance. The classification tree revealed that individuals displaying the maladaptive combination of low escape response speed, low boldness on the reef, and high routine swimming speed were highly susceptible to predation (92.4% with this combination died within 24 h). This accounted for 55.2% of all fish that died. Several combinations of behavioral traits predicted likely survival over 24 h, but there was greater uncertainty about that prediction than there was for fish that were predicted to die. Thus maladaptive behavioral traits were easier to identify than adaptive traits.     

Exposure to predation generates personality in threespined sticklebacks (Gasterosteus aculeatus)

A perplexing new question that has emerged from the recent surge of interest in behavioural syndromes or animal personalities is – why do individual animals behave consistently when behavioural flexibility is advantageous? If individuals have a tendency to be generally aggressive, then a relatively aggressive individual might be overly aggressive towards offspring, mates or even predators. Despite these costs, studies in several taxa have shown that individuals that are more aggressive are also relatively bold. However, the behavioural correlation is not universal; even within a species, population comparisons have shown that boldness and aggressiveness are correlated in populations of sticklebacks that are under strong predation pressure, but not in low predation populations. Here, we provide the first demonstration that an environmental factor can induce a correlation between boldness and aggressiveness. Boldness under predation risk and aggressiveness towards a conspecific were measured before and after sticklebacks were exposed to predation by trout, which predated half the sticklebacks. Exposure to predation generated the boldness–aggressiveness behavioural correlation. The behavioural correlation was produced by both selection by predators and behavioural plasticity. These results support the hypothesis that certain correlations between behaviours might be adaptive in some environments.     

Picking personalities apart: estimating the influence of predation,sex and body size on boldness in the guppy Poecilia reticulata

Predation is a strong selective force in most natural systems, potentially fueling evolutionary changes in prey morphology, life history and behaviour. Recent work has suggested that contrasting predation pressures may lead to population differentiation in personality traits. However, there are indications that these personality traits also differ between sexes and not necessarily in a consistent way between populations. We used an integrative approach to quantify boldness (latency to emerge from a shelter) in wild‐caught guppies in relation to predation pressure, population origin, sex and size. In addition we quantified the repeatability of these personality traits. We show that predation regime had significant effects on emergence time. In general, fish from high predation localities emerged sooner from the shelter compared to those from low predation localities. We found strong sex differences; males were significantly bolder than females. The relationship between emergence time and body size was non‐significant in all populations. We discuss what responses to expect from predator‐naïve versus predator‐experienced individuals and how this can be linked to the shyness–boldness continuum.     

Growth-mortality tradeoffs and 'personality traits' in animals

Consistent individual differences in boldness, reactivity, aggressiveness, and other 'personality traits' in animals are stable within individuals but vary across individuals, for reasons which are currently obscure. Here, I suggest that consistent individual differences in growth rates encourage consistent individual differences in behavior patterns that contribute to growth-mortality tradeoffs. This hypothesis predicts that behavior patterns that increase both growth and mortality rates (e.g. foraging under predation risk, aggressive defense of feeding territories) will be positively correlated with one another across individuals, that selection for high growth rates will increase mean levels of potentially risky behavior across populations, and that within populations, faster-growing individuals will take more risks in foraging contexts than slower-growing individuals. Tentative empirical support for these predictions suggests that a growth-mortality perspective may help explain some of the consistent individual differences in behavioral traits that have been reported in fish, amphibians, reptiles, and other animals with indeterminate growth.     

Variation in aggressive behaviour in the poeciliid fish Brachyrhaphis episcopi: Population and sex differences

Aggression is often positively correlated with other behavioural traits such as boldness and activity levels. Comparisons across populations can help to determine factors that promote the evolution of such traits. We quantified these behaviours by testing the responses of wild-caught poeciliid fish, Brachyrhaphis episcopi, to mirror image stimuli. This species occurs in populations that experience either high or low levels of predation pressure. Previous studies have shown that B. episcopi from low predation environments are less bold than those that occur with many predators. We therefore predicted that fish from high predation populations would be more aggressive and more active than fish from low predation populations. However, we found the opposite - low predation fish approached a mirror and a novel object more frequently than high predation fish suggesting that ‘boldness’ and aggression were higher in low predation populations, and that population-level boldness measures may vary depending on context. When tested individually, low predation fish inspected their mirror image more frequently. Females, but not males, from low predation sites were also more aggressive towards their mirror image. Variation in female aggression may be driven by a trade-off between food availability and predation risk. This suggests that the relationship between aggression and boldness has been shaped by adaptation to environmental conditions, and not genetic constraints.     

Two experimental designs generate contrasting patterns of behavioral differentiation along a latitudinal gradient in Lestes sponsa—Common‐garden not so common after all?

Understanding why and how behavioral profiles differ across latitudes can help predict behavioral responses to environmental change. The first response to environmental change that an organism exhibits is commonly a behavioral response. Change in one behavior usually results in shifts in other correlated behaviors, which may adaptively or maladaptively vary across environments and/or time. However, one important aspect that is often neglected when studying behavioral expressions among populations is if/how the experimental design might affect the results. This is unfortunate since animals often plastically modify their behavior to the environment, for example, rearing conditions. We studied behavioral traits and trait correlations in larvae of a univoltine damselfly, Lestes sponsa, along its latitudinal distribution, spreading over 3,300 km. We compared behavioral profiles among larvae grown in two conditions: (a) native temperatures and photoperiods or (b) averaged constant temperatures and photoperiods (common‐garden). We hypothesized latitudinal differences in behavioral traits regardless of the conditions in which larvae were grown, with northern populations expressing higher activity, boldness, and foraging efficiency. When grown in native conditions, northern larvae were bolder, more active and more effective in prey capture than central and low latitude populations, respectively, as well as showed the strongest behavioral correlations. In contrast, larvae reared in common‐garden conditions showed no differences between regions in both individual traits and trait correlations. The results suggest different selective pressures acting on the studied traits across populations, with environment as a central determinant of the observed trait values. Common‐garden designed experiments may evoke population‐dependent levels of plastic response to the artificial conditions and, hence, generate results that lack ecological relevance when studying multi‐population differences in behavior.     

A multi-trait,field-based examination of personality in a semi-aquatic turtle

Studies documenting and quantifying personality traits are common in animal behavior. Such studies often consider the nature of individual variation and personality as correlated with a variety of natural history, physiological, or ecological traits, and therefore consider the importance of personality for strategies in wild systems. Though such studies have contributed markedly to our understanding of the important aspects of personality that may covary with a variety of factors that might affect fitness, much of the research on animal personalities is taxonomically limited. To supplement and compliment the laboratory study of reptile personality, we examined the patterns of multiple personality traits in wild Eastern painted turtles (Chrysemys picta) in the field. We examined patterns of aggression, sociability, and boldness across these traits in different contexts in 103 adult, wild-caught turtles. We found strong correlations both within and among the focal behavioral axes, representing robust evidence for personality in this species. Specifically, we found strong relationships among many measures of aggression, as well as relationships between aggression and our measures of boldness. Finally, we note a tendency for sociability in our turtles, with animals scoring high on sociability showing lower tendency toward aggression. Overall, our study provides robust evidence for correlated suites of behavioral traits, or personality, in a semi-aquatic turtle surveyed in the wild. Future work should continue to expand the range of traits examined to more fully consider the ecological consequences of variation in personality in this and similar species.     

Predation risk and abiotic habitat parameters affect personality traits in extremophile populations of a neotropical fish (Poecilia vivipara)

Understanding whether and how ambient ecological conditions affect the distribution of personality types within and among populations lies at the heart of research on animal personality. Several studies have focussed on only one agent of divergent selection (or driver of plastic changes in behavior), considering either predation risk or a single abiotic ecological factor. Here, we investigated how an array of abiotic and biotic environmental factors simultaneously shape population differences in boldness, activity in an open‐field test, and sociability/shoaling in the livebearing fish Poecilia vivipara from six ecologically different lagoons in southeastern Brazil. We evaluated the relative contributions of variation in predation risk, water transparency/visibility, salinity (ranging from oligo‐ to hypersaline), and dissolved oxygen. We also investigated the role played by environmental factors for the emergence, strength, and direction of behavioral correlations. Water transparency explained most of the behavioral variation, whereby fish from lagoons with low water transparency were significantly shyer, less active, and shoaled less than fish living under clear water conditions. When we tested additional wild‐caught fish from the same lagoons after acclimating them to homogeneous laboratory conditions, population differences were largely absent, pointing toward behavioral plasticity as a mechanism underlying the observed behavioral differences. Furthermore, we found correlations between personality traits (behavioral syndromes) to vary substantially in strength and direction among populations, with no obvious associations with ecological factors (including predation risk). Altogether, our results suggest that various habitat parameters simultaneously shape the distribution of personality types, with abiotic factors playing a vital (as yet underestimated) role. Furthermore, while predation is often thought to lead to the emergence of behavioral syndromes, our data do not support this assumption.     

A Continuum of Behavioral Plasticity in Urban and Desert Black Widows

Behavioral plasticity marks an individual's ability to modulate behavior across functional contexts. Behavioral syndromes, on the other hand, appear as consistent individual variation in behavior that is both repeatable for individuals within a functional context (e.g., consistent voracity toward prey) and correlated across contexts (e.g., high voracity toward prey and high levels of boldness toward enemies). Thus, adaptive plasticity and syndromes represent two extremes of a behavioral plasticity continuum upon which most behavioral phenotypes fall. We tested for both adaptive plasticity and behavioral syndromes in the western black widow spider, Latrodectus hesperus. We measured behavior in three contexts: startle, startle + prey, and startle + mate, and found (1) classic behaviorally plastic responses to predation risk, (2) high repeatability of behavior within contexts, and (3) evidence of a correlation between startle + prey and startle + mate contexts, indicative of a behavioral syndrome. As relative behavioral plasticity may vary across populations, we also compared urban and desert populations to test whether spiders from these habitats exhibit different behaviors and/or behavioral syndromes. While we found that urban males used in mating trials courted urban females significantly more than desert females, we found no other differences in the behavior of urban and desert black widows. Thus, black widows, regardless of habitat, are characterized by both context‐specific behavioral plasticity and across‐context correlations, presenting a phenotypic complexity that is likely exhibited, to varying degrees, by most organisms.     

Differences in aggression,activity and boldness between native and introduced populations of an invasive crayfish

Aggressiveness, along with foraging voracity and boldness, are key behavioral mechanisms underlying the competitive displacement and invasion success of exotic species. However, do aggressiveness, voracity and boldness of the invader depend on the presence of an ecologically similar native competitor in the invaded community? We conducted four behavioral assays to compare aggression, foraging voracity, threat response and boldness to forage under predation risk of multiple populations of exotic signal crayfish Pacifastacus leniusculus across its native and invaded range with and without a native congener, the Shasta crayfish P. fortis. We predicted that signal crayfish from the invaded range and sympatric with a native congener (IRS) should be more aggressive to outcompete a close competitor than populations from the native range (NR) or invaded range and allopatric to a native congener (IRA). Furthermore, we predicted that IRS populations of signal crayfish should be more voracious, but less bold to forage under predation risk since native predators and prey likely possess appropriate behavioral responses to the invader. Contrary to our predictions, results indicated that IRA signal crayfish were more aggressive towards conspecifics and more voracious and active foragers, yet also bolder to forage under predation risk in comparison to NR and IRS populations, which did not differ in behavior. Higher aggression/voracity/boldness was positively correlated with prey consumption rates, and hence potential impacts on prey. We suggest that the positive correlations between aggression/voracity/boldness are the result of an overall aggression syndrome. Results of stream surveys indicated that IRA streams have significantly lower prey biomass than in IRS streams, which may drive invading signal crayfish to be more aggressive/voracious/bold to acquire resources to establish a population.     

Personality traits and behavioral syndromes in differently urbanized populations of house sparrows (Passer domesticus)

Urbanization creates novel environments for wild animals where selection pressures may differ drastically from those in natural habitats. Adaptation to urban life involves changes in various traits, including behavior. Behavioral traits often vary consistently among individuals, and these so-called personality traits can be correlated with each other, forming behavioral syndromes. Despite their adaptive significance and potential to act as constraints, little is known about the role of animal personality and behavioral syndromes in animals' adaptation to urban habitats. In this study we tested whether differently urbanized habitats select for different personalities and behavioral syndromes by altering the population mean, inter-individual variability, and correlations of personality traits. We captured house sparrows (Passer domesticus) from four different populations along the gradient of urbanization and assessed their behavior in standardized test situations. We found individual consistency in neophobia, risk taking, and activity, constituting three personality axes. On the one hand, urbanization did not consistently affect the mean and variance of these traits, although there were significant differences between some of the populations in food neophobia and risk taking (both in means and variances). On the other hand, both urban and rural birds exhibited a behavioral syndrome including object neophobia, risk taking and activity, whereas food neophobia was part of the syndrome only in rural birds. These results indicate that there are population differences in certain aspects of personality in house sparrows, some of which may be related to habitat urbanization. Our findings suggest that urbanization and/or other population-level habitat differences may not only influence the expression of personality traits but also alter their inter-individual variability and the relationships among them, changing the structure of behavioral syndromes.     

Weevil x Insecticide: Does ‘Personality’ Matter?

An insect’s behavior is the expression of its integrated physiology in response to external and internal stimuli, turning insect behavior into a potential determinant of insecticide exposure. Behavioral traits may therefore influence insecticide efficacy against insects, compromising the validity of standard bioassays of insecticide activity, which are fundamentally based on lethality alone. By extension, insect ‘personality’ (i.e., an individual’s integrated set of behavioral tendencies that is inferred from multiple empirical measures) may also be an important determinant of insecticide exposure and activity. This has yet to be considered because the behavioral studies involving insects and insecticides focus on populations rather than on individuals. Even among studies of animal ‘personality’, the relative contributions of individual and population variation are usually neglected. Here, we assessed behavioral traits (within the categories: activity, boldness/shyness, and exploration/avoidance) of individuals from 15 populations of the maize weevil (Sitophilus zeamais), an important stored-grain pest with serious problems of insecticide resistance, and correlated the behavioral responses with the activity of the insecticide deltamethrin. This analysis was performed at both the population and individual levels. There was significant variation in weevil ‘personality’ among individuals and populations, but variation among individuals within populations accounted for most of the observed variation (92.57%). This result emphasizes the importance of individual variation in behavioral and ‘personality’ studies. When the behavioral traits assessed were correlated with median lethal time (LT₅₀) at the population level and with the survival time under insecticide exposure, activity traits, particularly the distance walked, significantly increased survival time. Therefore, behavioral traits are important components of insecticide efficacy, and individual variation should be considered in such studies. This is so because population differences provided only crude approximation of the individual personality in a restrained experimental setting likely to restrict individual behavior favoring the transposition of the individual variation to the population.     

Behavioral Syndromes Break Down in Urban Song Sparrow Populations

Animals in urban habitats face a number of unique stresses, including the necessity of dealing with high levels of human activity. Growing research suggests that: (1) inherent traits, as opposed to learned behavior, influence which species invade urban habitats, and (2) individuals exhibit behavioral syndromes that limit behavioral flexibility. As a result, perhaps only animals with inherently bold personalities successfully settle in areas of high human activity, and such animals may also exhibit correlated variation in other behavioral traits, such as territorial aggression. In this study, we examine boldness and aggression in several urban and rural populations of song sparrows (Melospiza melodia). We found that urban birds were both bolder toward humans and also showed higher levels of aggression. We found a correlation between boldness and aggression in all populations combined, but no correlation within urban populations. Our results agree with other recent studies of song sparrow behavior, suggesting that greater boldness and aggression are general features of urban song sparrow populations, and a lack of a correlation between boldness and aggression in urban habitats is a general phenomenon as well. Urban habitats may select for bold and aggressive birds, and yet the traits can vary independently. These results add to a small number of studies which find that behavioral syndromes break down in potentially high quality habitats.     

Repeatability and correlation of physiological traits: Do ectotherms have a “thermal type”?

Across a range of taxa, individuals within a species differ in suites of correlated traits. These trait complexes, known as syndromes, can have dramatic evolutionary consequences as they do not evolve independently but rather as a unit. Current research focuses primarily on syndromes relating to aspects of behavior and life history. What is less clear is whether physiological traits also form a syndrome. We measured 10 thermal traits in the delicate skink, Lampropholis delicata, to test this idea. Repeatability was calculated and their across‐context correlations evaluated. Our results were in alignment with our predictions in that individual thermal traits varied consistently and were structured into a physiological syndrome, which we are referring to as the thermal behavior syndrome (TBS). Within this syndrome, lizards exhibited a “thermal type” with each being ranked along a cold–hot continuum. Hot types had faster sprint speeds and higher preferred body temperatures, whereas the opposite was true for cold types. We conclude that physiological traits may evolve as a single unit driven by the need to maintain optimal temperatures that enable fitness‐related behaviors to be maximized.     

Behavioural Syndromes in Urban and Rural Populations of Song Sparrows

Animals in urban habitats are often noticeably bold in the presence of humans. Such boldness may arise due to habituation, as urban animals learn, through repeated exposure, that passing humans do not represent a threat. However, there is growing research suggesting that: (1) inherent traits, as opposed to learned behaviour, influence which species invade urban habitats, and (2) individuals exhibit individual personality traits that limit behavioural flexibility, with the possible result that not all individuals would be able to demonstrate an appropriate level of boldness in urban environments. As a result, perhaps only birds with inherently bold personalities could successfully settle in an area of high human disturbance, and further, we might also expect to see the existence of behavioural syndromes, where boldness is correlated with variation in other behavioural traits such as aggression. In this study, we examined boldness and territorial aggression in urban and rural populations of song sparrows. We found that urban birds were bolder towards humans and that urban birds also showed higher levels of territorial aggression. We also found an overall correlation between boldness and territorial aggression, suggesting that urban boldness may be part of a behavioural syndrome. However, we see no correlation between boldness and aggression in the urban population, and thus, more work is needed to determine the mechanisms accounting for high levels of boldness and aggression urban song sparrows.     

Environmental and genetic effects shape the development of personality traits in the mangrove killifish Kryptolebias marmoratus

Consistent individual differences in behaviour are well documented, for example, individuals can be defined as consistently bold or consistently shy. To date our understanding of the mechanisms underpinning consistent individual differences in behaviour (also termed behavioural types (BTs)) remains limited. Theoretical work suggests life‐history tradeoffs drive BT variation, however, empirical support is scarce. Moreover, whilst life‐history is known to be phenotypically plastic in response to environmental conditions during ontogeny, the extent to which such plasticity drives plasticity in behavioural traits and personality remains poorly understood. Using a natural clonal vertebrate, Kryptolebias marmoratus, we control for genetic variation and investigate developmental plasticity in life‐history and three commonly studied behavioural traits (exploration, boldness, aggression) in response to three ecologically relevant environments; conspecific presence, low food and perceived risk. Simulated predation risk was the only treatment that generated repeatable behaviour i.e. personality during ontogeny. Treatments differed in their effects on mean life‐history and behavioural scores. Specifically, low food fish exhibited reduced growth rate and exploration but did not differ from control fish in their boldness or aggression scores. Conspecific presence resulted in a strong negative effect on mean aggression, boldness and exploration during ontogeny but had minimal effect on life‐history traits. Simulated predation risk resulted in increased reproductive output but had minimal effect upon average behavioural scores. Together these results suggest that life‐history plasticity/variation may be insufficient in driving variation in personality during development. Finally, using offspring derived from each rearing environment we investigate maternal effects and find strong maternal influence upon offspring size, but not behaviour. These results highlight and support the current understanding that risk perception is important in shaping personality, and that social experience during ontogeny is a major influence upon behavioural expression.     

Explaining leptokurtic movement distributions: intrapopulation variation in boldness and exploration

Leptokurtic distributions of movement distances observed in field-release studies, in which some individuals move long distances while most remain at or near their release point, are a common feature of mobile animals. However, because leptokurtosis is predicted to be transient in homogeneous populations, persistent leptokurtosis suggests a population heterogeneity. We found evidence for a heterogeneity that may generate persistent leptokurtosis. We tested individuals of the Trinidad killifish Rivulus hartii for boldness in a tank test and released them back into their native stream. Boldness in the tank test predicted distance moved in the field releases, even after effects of size and sex were removed. Further, data from a 19-mo mark-recapture study showed that individual growth correlated positively with movement in a predator-threatened river zone where the Rivulus population is spatially fragmented and dispersal is likely to be a hazardous activity. In contrast, no such correlation existed in a predator-absent zone where the population is unfragmented. These results show that a behavioral trait, not discernible from body size or sex, contributes to dispersal and that a component of fitness of surviving "dispersers" is elevated above that of "stayers," a fundamental assumption or prediction of many models of the evolution of dispersal through hazardous habitat.     

Linking behavioral thermoregulation,boldness, and individual state in male Carpetan rock lizards

Mechanisms affecting consistent interindividual behavioral variation (i.e., animal personality) are of wide scientific interest. In poikilotherms, ambient temperature is one of the most important environmental factors with a direct link to a variety of fitness‐related traits. Recent empirical evidence suggests that individual differences in boldness are linked to behavioral thermoregulation strategy in heliothermic species, as individuals are regularly exposed to predators during basking. Here, we tested for links between behavioral thermoregulation strategy, boldness, and individual state in adult males of the high‐mountain Carpetan rock lizard (Iberolacerta cyreni). Principal component analysis revealed the following latent links in our data: (i) a positive relationship of activity with relative limb length and color brightness (PC1, 23% variation explained), (ii) a negative relationship of thermoregulatory precision with parasite load and risk‐taking (PC2, 20.98% variation explained), and (iii) a negative relationship between preferred body temperature and relative limb length (PC3, 19.23% variation explained). We conclude that differences in boldness and behavioral thermoregulatory strategy could be explained by both stable and labile state variables. The moderate link between behavioral thermoregulatory strategy and risk‐taking personality in our system is plausibly the result of differences in reproductive state of individuals or variation in ecological conditions during the breeding season.     

Behavioral types of predator and prey jointly determine prey survival: potential implications for the maintenance of within-species behavioral variation

Recent studies in animal behavior have emphasized the ecological importance of individual variation in behavioral types (e.g., boldness, activity). Such studies have emphasized how variation in one species affects its interaction with other species. But few (if any) studies simultaneously examine variation in multiple interacting species, despite the potential for coevolutionary responses to work to either maintain or eliminate variation in interacting populations. Here, we investigate how individual differences in behavioral types of both predators (ocher sea stars, Pisaster ochraceus) and prey (black turban snails, Chlorostoma funebralis) interact to mediate predation rates. We assessed activity level, degree of predator avoidance behavior, and maximum shell diameter of individual C. funebralis and activity levels of individual P. ochraceus. We then placed 46 individually marked C. funebralis into outdoor mesocosms with a single P. ochraceus and allowed them to interact for 14 days. Overall, predator avoidance behavior and maximum shell diameter were positively associated with survival for C. funebralis. However, the effects of these traits depended on the predator's behavioral type: greater predator avoidance behavior was favored with active P. ochraceus, and low predator avoidance behavior was favored with inactive P. ochraceus. We argue that, even in two-species interactions, trait variation in heterospecifics could be an important factor maintaining trait variation within populations.