Mechanisms of benthic prey capture in wrasses (Labridae)

Teleost fishes capture prey using ram, suction, and biting behaviors. The relative use of these behaviors in feeding on midwater prey is well studied, but few attempts have been made to determine how benthic prey are captured. This issue was addressed in the wrasses (Labridae), a trophically diverse lineage of marine reef fishes that feed extensively on prey that take refuge in the benthos. Most species possess strong jaws with stout conical teeth that appear well-suited to gripping prey. Mechanisms of prey capture were evaluated in five species encompassing a diversity of feeding ecologies: Choerodon anchorago (Bloch, 1791), Coris gaimard (Quoy and Gaimard, 1824), Hologymnosus doliatus (Lacepède, 1801), Novaculichthys taeniourus (Lacepède, 1801) and Oxycheilinus digrammus (Lacepède, 1801). Prey capture sequences were filmed with high-speed video at the Lizard Island Field Station (14°40′S, 145°28′E) during April and May 1998. Recordings were made of feeding on pieces of prawn suspended in the midwater and similar pieces of prawn held in a clip that was fixed to the substratum. Variation was quantified among species and between prey types for kinematic variables describing the magnitude and timing of jaw, hyoid, and head motion. Species differed in prey capture kinematics with mean values of most variables ranging between two and four-fold among species and angular velocity of the opening jaw differing seven-fold. The kinematics of attached prey feeding could be differentiated from that of midwater captures on the basis of faster angular velocities of the jaws and smaller movements of cranial structures which were of shorter duration. All five species used ram and suction in combination during the capture of midwater prey. Surprisingly, ram and suction also dominated feedings on attached prey, with only one species making greater use of biting than suction to remove attached prey. These data suggest an important role for suction in the capture of benthic prey by wrasses. Trade-offs in skull design associated with suction and biting may be particularly relevant to understanding the evolution of feeding mechanisms in this group. Published online: 11 July 2002     

Wait before running for your life: defensive tactics of spiny mice (<Emphasis Type="Italic">Acomys cahirinus</Emphasis>) in evading barn owl (<Emphasis Type="Italic">Tyto alba</Emphasis>) attack

Raptor–prey encounters were studied to evaluate the strategies and success rate of both predator attack and prey defense. We compared the success of barn owls in catching stationary simulated prey (food item) with that of moving prey (food item that was pulled in various directions). We also tracked real encounters between barn owls and spiny mice in a captive environment. It was found that owls had higher success in attacking stationary prey and that they seemed to attack the prey as soon as it became motionless. When attacked, only a few spiny mice remained immobile (freeze response) whereas most fled and usually avoided capture by the owls. It was also found that spiny mice displayed a preference to escape in those directions in which owls had demonstrated a lower success in catching the simulated prey. Escape initiation dichotomized to a short or long (but rarely intermediate) distance between the spiny mouse and the owl with more successful avoidance at short-distance (last-moment) escapes. The best predictor of escape success was the velocity of the spiny mouse, and the second best predictor was its flight initiation distance (FID). We present an update for Ydenberg and Dill’s model for optimal FID in close encounters, suggesting that fleeing at the last moment is advantageous. However, a last-moment attempt to escape is also more risky with a split second differing between life and death, and is therefore appropriate mainly for agile prey under close-distance attack.     

Effects of water motion and prey behavior on zooplankton capture by two coral reef fishes

Water motion is an important factor affecting planktivory on coral reefs. The feeding behavior of two species of tube-dwelling coral reef fish (Chaenopsidae) was studied in still and turbulent water. One species of blenny, Acanthemblemaria spinosa , lives in holes higher above the reef surface and feeds mainly on calanoid copepods, while a second, A. aspera , lives closer to the reef surface, feeds mainly on harpacticoid copepods, and is exposed to less water motion than the first. In the laboratory, these two blenny species were video recorded attacking a calanoid copepod ( Acartia tonsa, evasive prey) and an anostracan branchiopod (nauplii of Artemia sp., passive prey). Whereas A. spinosa attacked with the same vigor in still and turbulent water, A. aspera modulated its attack with a more deliberate strike under still conditions than turbulent conditions. For both fish species combined, mean capture success when feeding on Artemia sp. was 100% in still water and dropped to 78% in turbulent water. In contrast, when feeding on Acartia tonsa, mean capture success was 21% in still water and rose to 56% in turbulent water. We hypothesize that, although turbulence reduces capture success by adding erratic movement to Artemia sp. (passive prey), it increases capture success of Acartia tonsa (evasive prey) by interfering with the hydrodynamic sensing of the approaching predator. These opposite effects of water motion increase the complexity of the predator-prey relationship as water motion varies spatially and temporally on structurally complex coral reefs. Some observations were consistent with A. aspera living in a lower energy benthic boundary layer as compared with A. spinosa: slower initial approach to prey, attack speeds modulated according to water velocity, and lower proportion of approaches that result in strikes in turbulent water.Communicated by P.W. Sammarco, Chauvin     

Morphology and kinematics of prey capture in the syngnathid fishes Hippocampus erectus and Syngnathus floridae

The mechanism of prey capture in two syngnathid fishes, the lined seahorse Hippocampus erectus (Perry) and the dusky pipefish Syngnathus floridae (Jordan and Gilbert), is described based on anatomical observations and high-speed video recordings (200 and 400 images s⁻¹) of feeding events by four seahorses and three pipefish. The fish were collected near Turkey Point, Florida, U.S.A., in January 1994 to March 1995. The dominant features of the morphology of these and many other syngnathiform fishes include extreme elongation of the suspensorium and neurocranium with a small mouth located at the anterior tip of the head. In the seahorse, a preparatory phase of prey capture consisted of slow ventral head flexion. This was followed by rapid elevation of the head and snout as the prey was drawn into the mouth by suction. Both H. erectus and S. floridae capture prey rapidly, with peak head excursions and mouth opening occurring within 5 to 7 ms of the onset of the strike. There was no upper jaw protrusion. In both species the time to recovery of the cranium and hyoid apparatus to resting positions was highly variable but took at least 500 ms. Manipulations of freshly dead specimens indicated a biomechanical linkage between head elevation and hyoid depression. However, the predictions of a previously proposed four-bar linkage model that couples hyoid depression to head elevation were not fully supported by kinematic data from one seahorse, suggesting that additional linkages act during the expansive phase of prey capture. These species exhibit the generalized kinematic pattern of prey capture in bony fishes, with head elevation, hyoid depression and mouth opening occurring almost simultaneously. The derived morphology results in a unique feeding behav‐ior, in which prey are captured during a sudden up-swing of the head, which brings the mouth to the prey. Suction is used to draw the prey into the buccal cavity. Received: 4 August 1996 / Accepted: 27 August 1996     

Precopulatory sexual cannibalism in fishing spiders (Dolomedes triton): a role for behavioral syndromes

Precopulatory sexual cannibalism (predation of a potential mate prior to copulation) offers an extreme example of intersexual conflict, a current focus in behavioral ecology. The aggressive-spillover hypothesis, posits that precopulatory sexual cannibalism may be a nonadaptive by-product of a general syndrome of voracity (aggression towards prey) that is expressed in multiple behavioral contexts. In this view, selection favoring high levels of voracity throughout ontogeny spills over to cause sexual cannibalism in adult females even when it is not necessarily beneficial. Using the North American fishing spider, Dolomedes triton, we present the first in depth test of this hypothesis. We found support for three aspects of the spillover hypothesis. First, voracity towards hetero-specific prey results in high feeding rates, large adult size, and increased fecundity. Second, juvenile and adult voracity are positively correlated (i.e., voracity is a consistent trait over ontogeny). Third, voracity towards hetero-specific prey is indeed positively correlated with precopulatory sexual cannibalism. Assays of antipredator behavior further revealed positive correlations between boldness towards predators, voracity and precopulatory sexual cannibalism. Overall, our results support the notion that precopulatory sexual cannibalism in D. triton is part of a behavioral syndrome spanning at least three major contexts: foraging, predator avoidance, and mating.     

The effects of zooplankton swimming behavior on prey-capture kinematics of red drum larvae, <Emphasis Type="Italic">Sciaenops ocellatus</Emphasis>

Most marine fishes undergo a pelagic larval phase, the early life history stage that is often associated with a high rate of mortality due to starvation and predation. We present the first study that examines the effects of prey swimming behavior on prey-capture kinematics in marine fish larvae. Using a digital high-speed video camera, we recorded the swimming velocity of zooplankton prey (Artemia franciscana, Brachionus rotundiformis, a ciliate species, and two species of copepods) and the feeding behavior of red drum (Sciaenops ocellatus) larvae. From the video recordings we measured: (1) zooplankton swimming velocity in the absence of a red drum larva; (2) zooplankton swimming velocity in the presence of a red drum larva; and (3) the excursion and timing of key kinematic events during prey capture in red drum larvae. Two-way ANOVA revealed that: (1) swimming velocity varied among zooplankton prey; and (2) all zooplankton prey, except rotifers and ciliates, increased their swimming velocity in the presence of a red drum larva. The kinematics of prey capture differed between two developmental stages in S. ocellatus larvae. Hyoid-stage larvae (3–14 days old) fed on slow swimming B. rotundiformis (rotifers) while hyoid-opercular stage larvae (15 days and older) ate fast moving A. franciscana. Hyoid-opercular stage red drum larvae had a larger gape, hyoid depression and lower jaw angle, and a longer gape cycle duration relative to their hyoid-stage conspecifics. Interestingly, the feeding repertoire within either stage of red drum development was not affected by prey type. Knowledge of the direct relationship between fish larvae and their prey aids in our understanding of optimal foraging strategies and of the sources of mortality in marine fish larvae.     

Coordinated mechanics of feeding,swimming, and eye movements in <Emphasis Type="Italic">Tautoga onitis</Emphasis>, and implications for the evolution of trophic strategies in fishes

Fish feeding behavior results from successful coordination of the fins, jaws, and sensory systems, and the organization of this behavior may affect the fish’s foraging abilities and trophic ecology. Using quantitative kinematic methods, movements of the jaws, fins and eyes of Tautoga onitis (Teleostei: Labridae) were analyzed during feeding events. Tautog feeding events consisted of three phases: approach, strike, and recovery, each defined by a combination of kinematic events. The approach was characterized by changes in fin movements and in body position, with the eyes directed forward at the food item. The strike began with the onset of jaw opening and protrusion, then cranial elevation, with the eyes no longer looking at the food item. The end of the strike and the beginning of the recovery involved a braking maneuver with the pectoral fins; the fish turns and swims away from the original food location item after prey capture. The coordination performance variables of tautog were quantitatively compared to published data from closely related cheiline wrasses and parrotfishes, which represent different feeding ecologies within a monophyletic assemblage. Fishes feeding on molluscs and benthic invertebrates (Cheilinus fasciatus and Tautoga onitis) represented an intermediate coordination condition, with herbivores (the parrotfishes, Scarus quoyi and Sparisoma radians) at one extreme, and fishes feeding on elusive prey (Epibulus insidiator and Oxycheilinus digrammus) at the other extreme. The analysis suggests that the biomechanical demands of coordination for feeding on benthic invertebrates may represent a generalized, and perhaps ancestral behavior in the wrasses, whereas more specialized trophic niches have evolved divergent, more specialized demands. Examining the movement and coordination of the jaws, fins, and eyes during fish feeding provides a detailed mechanistic basis for behavior, and comparison of coordination patterns during feeding among different taxa can measure how these trophic strategies differ. Understanding the evolution of feeding ecologies in these demersal fishes may have implications for understanding their role within their shallow water reef community.     

Feeding over the 24-h cycle: dietary flexibility of cathemeral collared lemurs (<Emphasis Type="Italic">Eulemur collaris</Emphasis>)

Animals show specific morphological, physiological and behavioural adaptations to diurnal or nocturnal activity. Cathemeral species, i.e. animals with activities distributed over the 24-h period, have to compromise between these specific adaptations. The driving evolutionary forces and the proximate costs and benefits of cathemerality are still poorly understood. Our goal was to evaluate the role of predator avoidance, food availability and diet quality in shaping cathemeral activity of arboreal mammals using a lemur species as an example. For this, two groups of collared lemurs, Eulemur collaris, were studied for 14 months in the littoral forest of southeastern Madagascar. Data on feeding behaviour were collected during all-day and all-night follows by direct observation. A phenological transect containing 78 plant species was established and monitored every 2 weeks to evaluate food availability during the study period. Characteristics of food items and animal nutritional intake were determined via biochemical analyses. The ratio of diurnal to nocturnal feeding was used as response variable in the analyses. The effects of abiotic environmental variables were removed statistically before the analyses of the biotic variables. We found that diurnal feeding lasted longer during the hot–wet season (December–February), whereas nocturnal feeding peaked during the hot–dry and cool–wet seasons (March–August). Although the lemurs foraged mostly in lower forest strata during daylight and used emergent trees preferably at night, the variables which measured animal exposure to birds of prey failed to predict the variation of the ratio of diurnal/nocturnal feeding. Ripe fruit availability and fiber intake are the two variables which best predicted the annual variation of the lemur diurnality. The data indicate that feeding over the whole 24-h cycle is advantageous during lean periods when animals have a fibre-rich, low-quality diet.     

Validity of the Prey‐Trap Hypothesis for Carnivore‐Ungulate Interactions at Wildlife‐Crossing Structures

Abstract: Wildlife‐exclusion fencing and wildlife‐crossing structures (e.g., underpasses and overpasses) are becoming increasingly common features of highway projects around the world. The prey‐trap hypothesis posits that predators exploit crossing structures to detect and capture prey. The hypothesis predicts that predation events occur closer to a highway after the construction of fences and crossing structures and that prey species’ use of crossings increases the probability that predators will attack prey. We examined interactions between ungulates and large carnivores at 28 wildlife crossing structures along 45 km of the Trans‐Canada Highway in Banff National Park, Alberta. We obtained long‐term records of locations where ungulates were killed (kill sites) before and after crossing structures were built. We also placed remote, motion‐triggered cameras at two crossing structures to monitor predator behavior following ungulate passage through the structure. The proximity of ungulate kill sites to the highway was similar before and after construction of fencing and crossing structures. We found only five kill sites near crossing structures after more than 32,000 visits over 13 years. We found no evidence that predator behavior at crossing structures is affected by prey movement. Our results suggest that interactions between large mammals and their prey at wildlife‐crossing structures in Banff National Park are not explained by the prey‐trap hypothesis.     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

In situ measures of foraging success and prey encounter reveal marine habitat-dependent search strategies

Predators are thought to reduce travel speed and increase turning rate in areas where resources are relatively more abundant, a behavior termed "area-restricted search." However, evidence for this is rare, and few empirical data exist for large predators. Animals exhibiting foraging site fidelity could also be spatially aware of suitable feeding areas based on prior experience; changes in movement patterns might therefore arise from the anticipation of higher prey density. We tested the hypothesis that regions of area-restricted search were associated with a higher number of daily speed spikes (a proxy for potential prey encounter rate) and foraging success in southern elephant seals (Mirounga leonina), a species exhibiting both area-restricted searches and high interannual foraging site fidelity. We used onshore morphological measurements and diving data from archival tags deployed during winter foraging trips. Foraging success was inferred from in situ changes in relative lipid content derived from measured changes in buoyancy, and first-passage time analysis was used to identify area-restricted search behavior. Seals exhibited relatively direct southerly movement on average, with intensive search behavior predominantly located at the distal end of tracks. The probability of being in search mode was positively related to changes in relative lipid content; thus, intensively searched areas were associated with the highest foraging success. However, there was high foraging success during the outward transit even though seals moved through quickly without slowing down and increasing turning rate to exploit these areas. In addition, the probability of being in search mode was negatively related to the number of daily speed spikes. These results suggest that movement patterns represent a response to prior expectation of the location of predictable and profitable resources. Shelf habitat was 4-9 times more profitable than the other habitats, emphasizing the importance of the East Antarctic shelf for this and other predators in the region. We have provided rare empirical data with which to investigate the relationship between predator foraging strategy and prey encounter/ foraging success, underlining the importance of inferring the timing and spatial arrangement of successful food acquisition for interpreting foraging strategies correctly.     

Feeding kinematics of hatchling swellsharks, Cephaloscyllium ventriosum (Scyliorhinidae): the importance of predator size

Capture, manipulation, and transport of prey were quantified from high-speed video of hatchling swellsharks, Cephaloscyllium ventriosum. Kinematic variables were contrasted with those of 1 yr-old swellsharks. Hatchling prey-captures were ram-dominated, while 1 yr-old prey-capture events had a detectable suction component. Timing differences between kinematic patterns of the age groups were not detected. Significant differences in displacement maxima of kinematic variables between the two age groups during feeding were detected, but were consistent with the expectations of isometry; they doubled in accordance with a doubling in shark length. A scaling analysis confirmed that swellsharks grow isometrically. A simple model of the head during prey capture confirmed that buccal expansion scaled isometrically between age groups. Thus, this study suggests that hatchlings generally perform the suite of movements necessary for suction generation within the buccal cavity during feeding. A suction component to the strike, however, was generally not detected by the “ram:suction index”. It appears that although it is probably generated within the buccal cavity, suction has little effect on the prey item and makes a minimal contribution to prey capture. Suction may be ineffective due to the highly active nature of the hatchlings. During a strike, a hatchling's forward locomotion may be sufficient to overwhelm any suction produced by the expanding buccal cavity; thus, the swimming shark effectively “scoops” the prey up in its open mouth (i.e. ram feeding) before the prey can be entrained in the flow of water entering the mouth (i.e. suction feeding). It is also likely that the hatchling sharks are sufficiently small to render any suction generated ineffective. Even though the sharks scale isometrically, the sheer size of the 1 yr-old sharks allows a greater amount of force to be generated, that will ultimately draw the prey to the open mouth. Thus, there are absolute consequences of size for feeding behaviors. Received: 17 June 1997 / Accepted: 6 March 1998     

Predator diversity and ecosystem functioning: density modifies the effect of resource partitioning

The link between biodiversity and ecosystem functioning is now well established, but the challenge remains to develop a mechanistic understanding of observed effects. Predator-prey interactions provide an opportunity to examine the role of resource partitioning, thought to be a principal mediator of biodiversity-function relationships. To date, interactions between multiple predators and their prey have typically been investigated in simplified agricultural systems with limited scope for resource partitioning. Thus there remains a dearth of studies examining the functional consequences of predator richness in diverse food webs. Here, we manipulated a species-rich intertidal food web, crossing predator diversity with total predator density, to simultaneously examine the independent and interactive effects of diversity and density on the efficiency of secondary resource capture. The effect of predator diversity was only detectable at high predator densities where competitive interactions between individual predators were magnified; the rate of resource capture within the species mixture more than doubled that of the best-performing single species. Direct observation of species-specific resource use in monoculture, as quantified by patterns of prey consumption, provided clear evidence that species occupied distinct functional niches, suggesting a mechanistic explanation of the observed diversity effect.     

Influence of flow speed on the functional response of a passive suspension feeder,the spionid polychaete <Emphasis Type="Italic">Polydora cornuta</Emphasis>

Passive suspension feeders rely on surrounding flow to deliver food particles to them. Therefore, the classic conception of functional response (feeding rate vs. food concentration) may require modification to account for flow speed as a second independent variable. I compared the functional response of Polydora cornuta at different velocities and determined whether food capture was proportional to particle flux (concentration × velocity). To understand feeding responses at a mechanistic level, I measured the functional responses in terms of contact and capture rates and determined particle retention efficiency. Experiments were run separately with two sizes of food particles, and with juvenile or adult worms. For both worm sizes and both particle sizes, capture rate in weak flow was directly related to concentration, but in strong flow it was constant. Worms were therefore unable to benefit from abundant food when in strong flow. The critical velocity at which the capture rate became constant was lower for adult worms than for juvenile worms, and it was lower for small particles than for large particles. Retention efficiency was constant among all treatments, and the results for contact rate were essentially the same as for capture rate. Therefore, the mechanics of particle contact must explain the effects of velocity on the functional response. Contact rate was not a constant proportion of particle flux; treatments with similar fluxes yielded different contact rates depending on the strength of flow. The results appeared to be caused by a velocity-induced behavioral change in appendage posture that affects contact rates: in moderate flow, worms form their feeding palps into helical coils, which they tighten as the velocity increases. I suggest this behavior constrains suspension feeding rates and the mechanical selection between particle sizes when worms are in strong flow, and that the effect changes with ontogeny. Because the results are consistent with patterns in measured growth rates of P. cornuta, I hypothesize that this influence of velocity on the functional response can constrain growth and population dynamics in this species.     

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.     

Boldness by habituation and social interactions: a model

Most studies of animal personality attribute personality to genetic traits. But a recent study by Magnhagen and Staffan (Behav Ecol Sociobiol 57:295–303, 2005) on young perch in small groups showed that boldness, a central personality trait, is also shaped by social interactions and by previous experience. The authors measured boldness by recording the duration that an individual spent near a predator and the speed with which it fed there. They found that duration near the predator increased over time and was higher the higher the average boldness of other group members. In addition, the feeding rate of shy individuals was reduced if other members of the same group were bold. The authors supposed that these behavioral dynamics were caused by genetic differences, social interactions, and habituation to the predator. However, they did not quantify exactly how this could happen. In the present study, we therefore use an agent-based model to investigate whether these three factors may explain the empirical findings. We choose an agent-based model because this type of model is especially suited to study the relation between behavior at an individual level and behavioral dynamics at a group level. In our model, individuals were either hiding in vegetation or feeding near a predator, whereby their behavior was affected by habituation and by two social mechanisms: social facilitation to approach the predator and competition over food. We show that even if we start the model with identical individuals, these three mechanisms were sufficient to reproduce the behavioral dynamics of the empirical study, including the consistent differences among individuals. Moreover, if we start the model with individuals that already differ in boldness, the behavioral dynamics produced remained the same. Our results indicate the importance of previous experience and social interactions when studying animal personality empirically.     

Diel variability of feeding activity in haddock (Melanogrammus aeglifinus) larvae in the East Shetland area, North Sea

Investigations of factors affecting feeding success in fish larvae require knowledge of the scales of variability of the feeding process itself and the indices used to assess this variability. In this study, we measured short-term (diel) variability in feeding rates of wild haddock (Melanogrammus aeglifinus) larvae four times per day during a 10-d cruise in the northern North Sea. Feeding activity was evaluated using indices of gut fullness, prey digestive state and biochemical measurements (tryptic enzyme activity). The gut fullness and the enzyme activity indices indicated moderate to high rates of food consumption throughout the cruise. Time series analysis of the three indices showed significant diel variability in all indices and enabled identification of significant lags between food uptake and peak digestive enzyme activity. The typical pattern of food consumption and digestion was characterized by maximal ingestion of prey early in the evening (19:00 hrs) and peak digestive enzyme activity at 01:00  hrs. The time scale over which enzyme activities reacted to prey ingestion was ca. 6 h, and is consistent with expectations from controlled laboratory experiments with other larval fish species. Significant diel variability in tryptic enzyme activity suggests that attempts to relate this measure of feeding success to other variables (e.g. food concentrations) should take care to accommodate natural cycles in feeding activity before making statistical comparisons. Received: 29 October 1998 / Accepted: 18 June 1999     

Risk vs. reward: how predators and prey respond to aging olfactory cues

Many animals use olfaction to find food and avoid predators, and must negotiate environments containing odors of varying compositions, strengths, and ages to distinguish useful cues from background noise. Temporal variation in odor cues (i.e., “freshness”) seems an obvious way that animals could distinguish cues, yet there is little experimental evidence for this phenomenon. Fresh cues provide a more reliable indicator of donor presence than aged cues, but we hypothesize that the benefits of responding to aged cues depend on whether the cue indicates the proximity of a predator or a potential meal. As prey cannot remain eternally risk averse in response to predator odor, we predict that antipredator responses should diminish as predator cues age. In contrast, animals searching for food should investigate aged prey cues if investigation costs are sufficiently low and the potential benefit (a meal) sufficiently high; thus, we predict that predators will maintain interest in aged prey cues. We tested these ideas using free-ranging rats (Rattus spp.) in two separate experiments; firstly assessing giving-up densities in the presence of predator odor, and secondly examining investigation rates of prey odors. As predicted, giving-up densities dropped once predator odor had aged, but investigation rates remained similar for aged and fresh prey odor. Thus, rats used temporal variation in odor cues to evaluate the cost–benefit relationship of responding to predator and prey odors. We suggest that the ecological significance of variable cue age needs more research and should be considered when interpreting behavioral responses to olfactory information.     

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.     

Individual- and population-level responses of a keystone predator to geographic variation in prey

Investigating how food supply regulates the behavior and population structure of predators remains a central focus of population and community ecology. These responses will determine the strength of bottom-up processes through the food web, which can potentially lead to coupled top-down regulation of local communities. However, characterizing the bottom-up effects of prey is difficult in the case of generalist predators and particularly with predators that have large dispersal scales, attributes that characterize most marine top predators. Here we use long-term data on mussel, barnacle, limpet, and other adult prey abundance and recruitment at sites spread over 970 km to investigate individual- and population-level responses of the keystone intertidal sunstar Heliaster helianthus on the coast of Chile. Our results show that this generalist predator responds to changes in the supply of an apparently preferred prey, the competitively dominant mussel Perumytilus purpuratus. Individual-level parameters (diet composition, per capita prey consumption, predator size) positively responded to increased mussel abundance and recruitment, whereas population-level parameters (density, biomass, size structure) did not respond to bottom-up prey variation among sites separated by a few kilometers. No other intertidal prey elicited positive individual predator responses in this species, even though a large number of other prey species was always included in the diet. Moreover, examining predator-prey correlations at approximately 80, 160, and 200 km did not change this pattern, suggesting that positive prey feedback could occur over even larger spatial scales or as a geographically unstructured process. Thus individual-level responses were not transferred to population changes over the range of spatial scales examined here, highlighting the need to examine community regulation processes over multiple spatial scales.