The avoidance response of fathead minnows to chemical alarm cues: understanding the effects of donor gender and breeding condition

Summary. All animals are vulnerable to predation at some point in their lives and consequently prey organisms often develop effective risk assessment systems. For many aquatic species predation risk assessment occurs through the use of olfactory cues, including predator odours and alarm cues from damaged or disturbed conspecifics. When aquatic species encounter conspecific alarm cues they may respond, or not, based on specific information including cue concentration, health and size of the conspecific donor and potentially the gender and breeding condition of the donor. Previous laboratory studies have demonstrated that fathead minnows (Pimephales promelas) fail to respond to the skin extracts of breeding male minnows. The purpose of the current study was to verify these early laboratory findings in the field as well as to further investigate the effect of female reproductive state and donor gender on the response of minnows to damage-release alarm cues. Our results indicate that male breeding condition has a significant effect on how minnows will respond to conspecific cues. Minnows showed avoidance of cues of female minnows and male minnows not in breeding condition, in comparison to cues of breeding male minnows and cues of male and female swordtails. Neither the gender of non-breeding minnows nor the reproductive state of female minnows influenced the avoidance of minnows to alarm cues.     

The paradox of risk assessment: comparing responses of fathead minnows to capture-released and diet-released alarm cues from two different predators

Summary. Many aquatic prey are known to use chemical alarm cues to assess their risk of predation. In fishes, such alarm cues can be released either through damage of the epidermis during a predatory attack (capture-released) or through release from the predator feces (diet-released). In our study, we compared the importance of capture- versus diet-released alarm cues in risk assessment by fathead minnows (Pimephales promelas) that were na?ve to fish predators. We utilized two different fish predators: a specialized piscivore, the northern pike (Esox lucius) and a generalist predator, the brook trout (Salvelinus fontinalis). Handling time of pike consuming minnows was much shorter than for trout consuming minnows, likely resulting in less epidermal damage to the minnows during attacks by pike. In accordance with this, minnows showed a less intense antipredator response to capture-released cues from pike than capture-released cues from trout. This represents a paradox in risk assessment for the minnows as they respond to the specialized piscivore, the more dangerous predator, with a less intense antipredator response. In contrast, the minnows showed a stronger antipredator response to the specialized piscivore than to the generalist when given diet cues. This work highlights the need for researchers to carefully consider the nature of the information available to prey in risk assessment.     

Graded recruitment and hunting strategies linked to prey weight and size in the ponerine ant Ectatomma ruidum

According to the weight and size of their prey, Ectatomma ruidum workers can employ different recruitment systems (solitary hunting, cooperative hunting and group hunting with recruitment) when mastering and retrieving prey items from short distances from the nest. Prey size determined the backwards entry typically adopted by this species, while prey weight determined the predatory strategy selected. After a common initial sequence (search for prey, detection, localization), predatory sequences varied in terms of the type of approach, the site of seizure, the reaction after stinging and the type of transport. Nevertheless, irrespective of prey weight and size, seizure was preferentially oriented towards the head and prey were always stung. Short-range recruitment and mass recruitment without trail laying were elicited by a large range of heavy prey (> 2.5 times the weight of an individual worker). According to the mortality risk associated with each prey, hunters exhibited a “prudent” stinging posture associated with an increase in the duration of the subsequent phase of waiting for prey immobilization. The overall time of capture was positively correlated with the weight of the prey. When collective hunting strategies were involved, E. ruidum colonies matched the number of recruited hunters to the size and weight of the prey. Compared to solitary hunting strategies, for short food–nest distances, this graded recruitment appeared to enhance the energetic benefits derived by this species from the use of recruitment systems: the higher the number of workers involved in the recruitment process, the greater the energetic benefits obtained. The exhibition or absence of trail laying behavior in the recruitment responses displayed by E. ruidum workers is discussed in relation to their involvement in scavenging or predatory behavior. Received: 27 June 1996 / Accepted after revision: 3 March 1997     

Making sense of predator scents: investigating the sophistication of predator assessment abilities of fathead minnows

According to the threat-sensitive predator avoidance hypothesis, selection favors prey that accurately assess the degree of threat posed by a predator and adjust their anti-predator response to match the level of risk. Many species of animals rely on chemical cues to estimate predation risk; however, the information content conveyed in these chemical signatures is not well understood. We tested the threat-sensitive predator avoidance hypothesis by determining the specificity of the information conveyed to prey in the chemical signature of their predator. We found that fathead minnows (Pimephales promelas) could determine the degree of threat posed by northern pike (Esox lucius) based on the concentration of chemical cues used. The proportion of minnows that exhibited an anti-predator response when exposed to a predator cue increased as the concentration of the pike cue used increased. More surprisingly, the prey could also distinguish large pike from small pike based on their odor alone. The minnows responded more intensely to cues of small pike than to cues of large pike. In this predator–prey system small pike likely represent a greater threat than large pike.Communicated by A. Mathis     

The relative roles of predation and sperm competition on the duration of the post-copulatory association between the sexes in the blue crab, Callinectes sapidus

In many species, post-copulatory mate guarding prevents other males from mating with the guarded female. In crabs, males stay with their mates to protect the female from predators because, in some species, mating occurs when she is soft and vulnerable after molting. I tested the relative roles of sperm competition and predation on the duration of the post-copulatory association in the blue crab, Callinectes sapidus. Unpaired females suffered greater predation mortality than paired females and males stayed with the female longer in the presence of predators than in their absence, suggesting that the post-copulatory association protects females during their vulnerable period. However, the association may also occur in blue crabs because of sperm competition since spermathecal contents of females in the field indicate that 12.4% mated twice. Females experimentally mated with two males contained both males ejaculates and each ejaculate had access to the unfertilized eggs, suggesting that the size of a male's ejaculate influences his fertilization rate in a multiply-mated female. Males stayed longest in response to a high risk of sperm competition. Longer post-copulatory associations allowed the first male's ejaculate to harden into a type of sperm plug, which limited the size of a second inseminator's ejaculate in a non-virgin female as compared with a virgin. Males passed larger ejaculates in the presence of rivals and when previous ejaculates were in the female spermathecae, another response to sperm competition. Larger ejaculates may need longer post-copulatory associations before a more effective sperm plug forms. Large males stayed with the female longer, which is consistent with their ability to pass larger ejaculates than small males and suggests that there may be costs to minimizing the duration of the post-copulatory association. In the field, associations last long enough to protect the female during her vulnerable phase and may ensure that the guarding male fertilizes the most eggs in the female, even if she remates. Thus, the post-copulatory association protects female blue crabs from additional inseminators as well as from predators. Received: 23 January 1996 / Accepted after revision: 9 November 1996     

Hatching with the enemy: Daphnia diapausing eggs hatch in the presence of fish kairomones

Summary. Infochemicals are known to play a key role in mediating predator-prey interactions, both in aquatic and terrestrial communities. However, state-dependent variation may exist in how effectively individuals can use this information, depending on genotype, life stage and experience. For our study, we used the predator-prey model system fish-waterflea Daphnia magna Straus (Cladocera, Daphniidae). Adult Daphnia use fish-derived infochemicals, so-called kairomones, as indicators of predation risk, and exhibit a spectrum of morphological, behavioural and life-history responses to the presence of fish kairomones. Here, we investigate whether diapausing eggs, an embryonic resting stage in the life cycle of D. magna, also use fish kairomones and tune their hatching to the risk of fish predation, as reported for diapausing stages of dinoflagellates. In two laboratory experiments, we studied hatching proportion and time until hatching of D. magna diapausing eggs in the absence and presence of fish kairomones. D. magna families differed significantly in their response to the presence of fish kairomones; some families reduced hatching proportion, whereas others increased it. Our results imply genotype-dependent differences in the hatching reactions to fish kairomones as observed for other traits in adult Daphnia.     

Non-adaptive adoptions of nestlings in the colonial lesser kestrel: proximate causes and fitness consequences

We examined the causes, costs and benefits of adoption in the altricial lesser kestrel Falco naumanni. Specifically, we tested the intergenerational conflict hypothesis, proposed to explain adoption in some birds. Adoptions involved 76% of the nests and 51% of the nestlings at a mean age of 25 days (12 days before fledging). Nest-switching nestlings were not in poorer body condition, more parasitized or younger than their siblings, and body condition and prey delivery rates of their parents did not differ from those of other parents. In the foster nest, adopted nestlings did not benefit from higher feeding rates or a prolongation of the nestling period. They did not have fewer nest-mates or achieve higher rank within the new brood. Thus, adopted nestlings did not improve their body condition and survival. Adult lesser kestrels seemed unable to finely discriminate beween their own and alien chicks. Foster parents bore the cost of an increase of prey delivery rates, although it did not affect their survival or subsequent reproductive performance. Therefore, our results do not support the intergenerational conflict hypothesis, and suggest that adoption in this species is non-adaptive. Traditionally, the lesser kestrel bred in cliffs where movement among nest-sites was restricted. Nowadays, about half of the colonies are in tiled roofs which facilitate nest-switching by nestlings. The high rate of adoptions may thus be explained as reproductive errors associated with the recent occupation of a new breeding habitat. Received: 3 May 1996 / Accepted after revision: 19 January 1997     

Algal blooms decrease care but increase egg survival in a fish with paternal care

Human-induced eutrophication, resulting in algal blooms and increased water turbidity, is an alarming problem in aquatic systems. Here, we experimentally tested the impact of algal turbidity on parental care, egg fanning, and time in the nest, in the sand goby, Pomatoschistus minutus, a fish with uniparental male care. We allowed males to care for their eggs in either clear water or water made turbid by planktonic algae. In the early brood cycle, males fanned their eggs less in turbid than in clear water, but this difference disappeared later. Despite decreased care, egg survival was higher in turbid conditions, indicating that early fanning may partly be redundant for egg survival and perhaps used more as courtship. Males also spent more time out of their nest in turbid water, perhaps as a means to encounter additional females under conditions of low visibility.     

Common goldeneyes adjust maternal effort in relation to prior brood success and not current brood size

Parental investment theory predicts that parental effort should be related to the reproductive value of the current brood. This depends on both the number of young and the survival prospects of each of them. Thus parents may provide more care to larger broods either because of (1) the direct effect of brood size per se on reproductive value (the “brood size” hypothesis) or because (2) past mortality, reflected in current brood size, predicts future mortality of the brood and hence its reproductive value (the “brood success” hypothesis). Earlier studies have not attempted to distinguish between these alternatives. We tested the hypotheses in the precocial, nidifugous common goldeneye Bucephala clangula, a species with uniparental female care. Maternal effort was measured as the time spent by the female in rearing the brood. We found that brood size itself is not associated with maternal effort, but that females modify their maternal effort according to the mortality already experienced by the brood, supporting the prediction of the brood success hypothesis. We also found that brood mortality varied considerably between broods and that previous mortality predicts future mortality within broods, basic assumptions of the brood success hypothesis. Received: 30 January 1996 / Accepted after revision: 27 October 1996     

Bright plumage in the magpie: does it increase or reduce the risk of predation?

Conspicuous color patterns in birds may lead to increased risk of predation. Alternatively, bright birds may be aposematic or unprofitable prey, which leads to decreased predation. During four autumns, I examined whether the black-and-white plumage of (stuffed) magpies Pica pica increases or reduces the risk of attack by migrating goshawks Accipiter gentilis. Attack risk was higher for normal magpies than for cryptic, brown magpies whether the mounts were placed near one another or far apart. However, the brown magpie may have been avoided because of its novelty. For magpies and cryptic jays Garrulus glandarius exposed together, the attack risk was similar. In 2 years, magpie and jay mounts were also exposed far apart. In 1994, with an invasion of migrating jays, attack risk was much higher for jays than for magpies. In 1995, with a normal density of jays, hawks attacked the magpies more often. The results demonstrate frequency-dependent prey selection by goshawks, which would influence any predation cost of bright plumage. The attacks on normally colored magpie mounts suggest that magpies are not aposematic. Trials with photographs and human observers indicated that normal magpies were somewhat easier to detect than jays. The plumage of the magpie possibly increases the risk of predation, but may be favoured by sexual or social selection. Received: 18 January 1996 / Accepted after revision: 6 October 1996     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

An estimate of female mate searching costs in the lekking butterfly Coenonympha pamphilus

We estimated the cost to females of the lekking butterfly Coenonympha pamphilus of visiting males on leks instead of taking off and soliciting courtship from males passing them outside leks, as occurs in the non-lekking congener C. tullia. We followed released virgin females of C. pamphilus in the field until they mated. We estimated the time they lost by remaining perched and not approaching males that passed them, and observed how often and at what distance virgin females were passed by males in the field. We then calculated how much faster these females would have been detected if they had taken off and approached these males, using probabilities of detection as a function of passing distance derived from field observations on C. tullia females. C. pamphilus females on average lost 201 min by not approaching males on their way to leks. To estimate what this time loss meant in terms of fitness, we measured how the age at mating affected fecundity. Using field measures of fecundity and mortality, the time loss translated into an average 2.8% reduction in fecundity as a best estimate, and an average 1.3% reduction in fecundity as a lower estimate. This fitness cost is larger than has been reported earlier for a lekking bird, but is probably too small to eliminate the possibility of indirect benefits of mating with males on leks. Received: 15 August 1996 / Accepted after revision: 24 January 1997     

Performance level and efficiency of two differing predator-avoidance strategies depend on nutritional state of the prey fish

Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.     

Consequences of patch reef spacing for density-dependent mortality of coral-reef fishes

The spatial configuration of habitat patches can profoundly affect a number of ecological interactions, including those between predators and prey. I examined the effects of reef spacing on predator-prey interactions within coral-reef fish assemblages in the Bahamas. Using manipulative field experiments, I determined that reef spacing influences whether and how density-dependent predation occurs. Mortality rates of juveniles of two ecologically dissimilar species (beaugregory damselfish and yellowhead wrasse) were similarly affected by reef spacing; for both species, mortality was density dependent on reef patches that were spatially isolated (separated by 50 m), and density independent on reef patches that were aggregated (separated by 5 m). A subsequent experiment with the damselfish demonstrated that a common resident predator (coney) caused a substantial proportion of the observed mortality, independent of reef spacing. Compared to isolated reefs, aggregated reefs were much more likely to be visited by transient predators (mostly yellowtail snappers), regardless of prey density, and on these reefs, mortality rates approached 100% for both prey species. Transient predators exhibited neither an aggregative response nor a type 3 functional response, and consequently were not the source of density dependence observed on the isolated reefs. These patterns suggest that resident predators caused density-dependent mortality in their prey through type 3 functional responses on all reefs, but on aggregated reefs, this density dependence was overwhelmed by high, density-independent mortality caused by transient predators. Thus, the spatial configuration of reef habitat affected both the magnitude of total predation and the existence of density-dependent mortality. The combined effects of the increasing fragmentation of coral reef habitats at numerous scales and global declines in predatory fish may have important consequences for the regulation of resident fish populations.     

Bold minnows consistently approach danger in the field and lab in response to either chemical or visual indicators of predation risk

A behavioral syndrome is a suite of behaviors correlated across multiple social contexts. In this study, boldness in the face of predation risk was assessed twice in fish across two different sensory modalities in both the field and lab to ascertain the biological relevance and complexity of this attribute. Individual fathead minnows were captured from a natural field population using traps that either contained chemical alarm cues (conspecific skin extract) or control (well water) and their responses to the presence of predator behind a glass partition assessed in the laboratory. Although fewer fish were captured in alarm cue-labeled traps, these bold fish performed longer predator inspections than shy fish captured in control traps. Thus, a shy/bold behavioral syndrome was expressed consistently across field and lab settings in response to both chemical and visual indicators of danger. Shy and bold individuals did not differ in sex, body length, secondary sexual characteristics, or parasite load but were of more robust physical condition.     

Why is competition more intense if food is supplied more slowly?

We investigated competition for food among two groups of six clone amagos (salmonids), Oncorhynchusmasoumacrostomus, in a laboratory experiment with different rates of food input. We examined the effect of temporal clumping of food resources on the inequality of food sharing between competitors. Monopolization of food by dominants was greater at a low input rate (one food item per 10 s) than at a high input rate (1 food item per 1 s). Aggressive behavior by dominants was more frequent at the low input rate than at the high input rate; its purpose was presumably to interfere with the feeding behavior of subordinates. We assessed the relative importance of three foraging factors (the number of approaches to food items, the chance per approach and the gain per chance) in enhancing inequality in food gain between individuals. Dominants had a disproportionately high chance per approach and gain per chance at the low input rate, but not at the high input rate. The chance of obtaining a food item per approach depended on how many competitors approached simultaneously. The gain per chance depended on the competitive ability of the approaching fish. There was an interaction between these components, such that the number of approaches affected the chance per approach and gain per chance. We evaluated the independent effect of the chance per approach, and showed that it was higher for dominants than for subordinates at the low input rate, but not at the high input rate. This implies that subordinates changed their behavior and became more likely to avoid approaching food at the same time as dominants at the low input rate. Received: 13 August 1996 / Accepted after revision: 30 November 1996     

Neighbours, strangers and male-male aggression as a determinant of lek size

Interactions between males on leks may play an influential role in lek formation and the regulation of lek size. In this paper I present the results of a playback experiment that simulated de novo settlement at sites adjacent to currently existing display territories of the ochre-bellied flycatcher, Mionectes oleagineus. In the study population, males displayed both solitarily and at small leks. A large proportion of males held no display territory at all. A stranger's song was played to both solitary and lekking males from 10 m outside their territorial boundaries. In separate playbacks, lekking males were also played neighbour's song. Both lekking and solitary territorial males reacted to the playback by decreasing their song rate, approaching the playback speaker and, on occasion, attacking the model. Solitarily displaying males responded more aggressively to playback of stranger's song than did lek males. Lek males were able to distinguish between their neighbour's and a stranger's song and did so irrespective of whether it was played from the neighbour's territory or from outside the lek. In addition to distinguishing between neighbours and strangers, lek males modified their responses to these different playbacks depending on where the playback originated. These results suggest that male-male interactions can be influential in structuring leks. In M. oleagineus, interactions between males are aggressive and act to limit rather than augment lek size. Received: 6 March 1996 / Accepted after revision: 9 December 1996     

Behavioral response races, predator-prey shell games, ecology of fear, and patch use of pumas and their ungulate prey

The predator-prey shell game predicts random movement of prey across the landscape, whereas the behavioral response race and landscape of fear models predict that there should be a negative relationship between the spatial distribution of a predator and its behaviorally active prey. Additionally, prey have imperfect information on the whereabouts of their predator, which the predator should incorporate in its patch use strategy. I used a one-predator-one-prey system, puma (Puma concolor)-mule deer (Odocoileus hemionus) to test the following predictions regarding predator-prey distribution and patch use by the predator. (1) Pumas will spend more time in high prey risk/low prey use habitat types, while deer will spend their time in low-risk habitats. Pumas should (2) select large forage patches more often, (3) remain in large patches longer, and (4) revisit individual large patches more often than individual smaller ones. I tested these predictions with an extensive telemetry data set collected over 16 years in a study area of patchy forested habitat. When active, pumas spent significantly less time in open areas of low intrinsic predation risk than did deer. Pumas used large patches more than expected, revisited individual large patches significantly more often than smaller ones, and stayed significantly longer in larger patches than in smaller ones. The results supported the prediction of a negative relationship in the spatial distribution of a predator and its prey and indicated that the predator is incorporating the prey's imperfect information about its presence. These results indicate a behavioral complexity on the landscape scale that can have far-reaching impacts on predator-prey interactions.     

The effect of predation on artificial reef juvenile demersal fish species

There is a concern that artificial reefs (AR) may act purely as fishing aggregation devices. Predators attracted to ARs can influence the distribution and abundance of prey fish species. Determining the role of predators in AR is important in advancing the understanding of community interactions. This paper documents the effects of predation on fish assemblages of AR located near a coastal lagoon fish nursery. The Dicentrarchus labrax is a very opportunistic species preying on juveniles (0⁺ and 1⁺ age classes) of several demersal fish species on the ARs. Reef prey and sea bass abundance were negatively correlated. The mean numbers of prey per sea bass stomach increased with the increase of reef fish prey abundance, suggesting that predation has a significant influence, resulting in a decrease in prey abundance. Prey mortality (4–48%) of demersal reef fish associated species depends on bass density. Prey selection was related both with prey abundance and vulnerability. Results showed that D. labrax predation on AR-fish associated species can increase prey natural mortality. However, the role of bass predation on the ecological functioning of exploited ARs is not clear. There may be increases in local fishing yields due either to an increase in predator biomass through aggregation of sea bass attracted to ARs or to greater production. In contrast, predation on juveniles of economically important reef fish preys, especially the most frequent and abundant (Boops boops), can contribute to a decrease in recruitment to the fishery. Our results indicate that inter-specific interactions (predator–prey) are important in terms of conservation and management, as well as for the evaluation of the long-term effects of reef deployment. Thus, it is necessary to consider ecological interactions, such as predation, prior to the development and deployment of artificial habitats as a tool for rehabilitation.     

The role of learning in the acquisition of threat-sensitive responses to predator odours

The supposition that prey animals respond to a predator with an intensity that matches the risk posed by the predator is known as the threat-sensitive predator avoidance hypothesis. Many studies have provided support for this hypothesis; yet, few studies have attempted to determine how such abilities are acquired by prey species. In this study, we investigated whether fathead minnows (Pimephales promelas) could learn to recognize an unknown predator (northern pike, Esox lucius) in such a way that they could match the intensity of their antipredator response with the threat posed by the predator. We exposed pike-naïve minnows to conspecific alarm cues paired with either a high or low concentration of pike odor. The following day, both groups were tested for a response to either high or low concentration of pike odor alone. We found that minnows conditioned with alarm cues paired with a given concentration of pike odor subsequently responded with a higher intensity to higher concentrations of pike odor, and with a lower intensity to lower concentrations of pike odor. These results demonstrate that during a single conditioning trial, minnows learn the identity of the predator in a threat-sensitive manner. Minnows use predator odor concentrations that they experience in subsequent interactions to adjust the intensity of their antipredator behavior.