How plankton copepods avoid fish predation: From individual responses to variations of the life cycle

Analysis of original and published data on predator avoidance by marine and freshwater plankton copepods, a major diet of many young fishes, suggests that individual defense mechanisms play a minor role in copepod anti-predator behavior. Capture success by planktivorous fish depends largely on prey visibility and the ability of the prey to escape. Copepods have almost no chance to avoid relatively large fish when encountered, but they can be ranked according to their ability to escape from larvae and fries. In contrast to small pelagic fish, which are also under heavy predation pressure, copepods rely more on prevention of the threat of predation than on active attempts to escape. Seeking a refuge in habitats non-accessible to predators would be more effective for these small and rather slow animals. Retreat into such refuges is accomplished by vertical and horizontal migrations, either diel, seasonal or ontogenetic. A decrease in activity (feeding, metabolism, reproduction, movement), resulting in diapause in deep water layers in its most pronounced version, is the ultimate attempt by copepods to separate from their predators, both in time and in space.     

Factors affecting feeding selectivity of visual predators on the copepod Acartia tonsa: locomotion,visibility and escape responses

Visual predation by fish on copepods involves prey encounter, attack and capture; during any of these processes prey selection can occur. Developmental changes in copepods, including increases in swimming speed, size and image contrast increase the encounter rate and distance at which they can be detected by predators. Copepods compensate for this increase vulnerability with age through diel vertical migration and improved escape capabilities. This study quantifies the changes in swimming speed and movement pattern with developmental stage of the copepod Acartia tonsa, using a video-computer system for motion analysis. Changes in visible size and image contrast with developmental stage were quantified under simulated natural illumination conditions using a video based image analysis system. The escape responses of the naupliar stages of the copepod Acartia tonsa were quantified in response to a stationary pipette sucking in water at a constant speed. Accurate quantification of the parameters that affect feeding selectivity of planktivorous fish will provide the basis for evaluation of their relative importance in future studies.     

The behavioural basis of prey selection by underyearling bream (Abramis brama (L.)) and roach (Rutilus rutilus (L.))

SUMMARY. A clear difference in the ability lo escape from fish predators exists between members of the Cladocera and Copepoda. The results of our laboratory studies have shown that underyearling roach and bream both found copepods more difficult to capture than cladocerans. However, bream were far more efficient than roach at catching the more elusive copepod prey. The basis for this difference was the greater strike ability of bream, most likely related to its more protrusible mouth. In their natural environment the two species of fish exhibited food resource partitioning with planktonic Cladocera predominating in roach guts and copepods and non-planktonic Cladocera composing the vast majority of the gut contents of bream. Differences in diet are partly due to their contrasting attack abilities. Additional variation may arise because of subtle differences in the timing and location of foraging.     

The Visually Mediated Escape Response in Fish: Predicting Prey Responsiveness and the Locomotor Behaviour of Predators and Prey

The outcome of predator-prey encounters is determined by a number of factors related to the locomotor and sensory performance of the animals. Escape responses can be triggered visually, i.e. by the magnifying retinal image of an approaching object (i.e. a predator), called the looming effect, and calculated as the rate of change of the angle subtended by the predator frontal profile as seen by the prey. A threshold of looming angle (ALT, the Apparent Looming Threshold) determines the reaction distance of a startled fish, which is proportional to the attack speed of the predator and its apparent frontal profile. Optimal tactics for predator attacks as well as consideration on their functional morphology are discussed in relation to ALT. Predator optimal attack speeds depend on predator morphology as well as the prey ALT. Predictions on the scaling of ALT suggest that ALT may increase (i.e. implying a decrease in reaction distance) with prey size in cases in which predator attack speeds are high (i.e. > 4 L/s in a 1-m long predator), while it may be relatively independent of prey size when predators attack at lower speeds. The issue of scaling of ALT is discussed using examples from field and laboratory studies. While the timing of the escape is a crucial issue for avoiding being preyed upon, the direction of escape manoeuvres may also determine the success of the escape. A simple theoretical framework for optimal escape trajectories is presented here and compared with existing data on escape trajectories of fish reacting to startling stimuli.     

An avoidance learning submodel for a general predation model

Summary This paper attempts to determine the effect on the number of prey eaten by predators of the addition of the component avoidance learning by prey to a computer model of the predation process developed by Holling. Generality was retained by concentrating upon a basic aspect of the prey's behaviour, its distance of reaction to an approaching predator. The zebra danio (Brachydanio rerio), a small freshwater fish, was used as an analogue of a general vertebrate prey. The predator used was the largemouth bass (Micropterus salmoides).Previous work (Dill, 1973b) showed that prey reactive distance increased with increasing experience with the predator. In the present study, this increased prey reactive distance is shown to increase predator pursuit time and hypothesized to decrease predator pursuit success. These relationships were expressed mathematically and built into Holling's (1965, 1966) model of the predation process, along with an equation describing the way in which reactive distance increases following an unsuccessful attack. Other changes necessitated in the model by the addition of the avoidance learning component included: a) Modifications of the calculation of search time to remove a previously implicit time spent unsuccessfully pursuing prey, and to correct the density of prey to account for those whose reactive distances exceed that of the predator and are therefore not susceptible to discovery; b) Addition of a new subroutine (CHASE) to calculate pursuit time, unsuccessful pursuit time, pursuit success, and strike success; c) Changes in subroutine ADCOM to assign prey to different classes (with different reactive distances) according to the number of times they have been unsuccessfully attacked; and d) Addition of a stochastic element via random numbers to determine the class to which an attacked prey belongs, the time to refuge, and the predator's strike success.Simulation was used to explore the consequences of these additions. The capability of learning substantially increased the prey's probability of surviving subsequent attack. Addition of an avoidance learning component caused declines in the predator's functional responses to both prey and predator density. The new component was also suggested to decrease the predator's numerical response to prey density and to increase the probability of stability in a predator-prey interaction.From a thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, University of British Columbia.     

Effects of successive predator attacks on prey aggregations

We study the cumulative effect of successive predator attacks on the disturbance of a prey aggregation using a modelling approach. Our model intends to represent fish schools attacked by both aerial and underwater predators. This individual-based model uses long-distance attraction and short-distance repulsion between prey, which leads to prey aggregation and swarming in the absence of predators. When intermediate-distance alignment is added to the model, the prey aggregation displays a cohesive displacement, i.e., schooling, instead of swarming. Including predators, i.e. with repulsion behaviour for prey to predators in the model, leads to flash expansion of the prey aggregation after a predator attack. When several predators attack successively, the prey aggregation dynamics is a succession of expanding-grouping-swarming/schooling phases. We quantify this dynamics by recording the changes in the simulated prey aggregation radius over time. This radius is computed as the longest distance of individual prey to the aggregation centroid, and it is assumed to increase along with prey disturbance. The prey aggregation radius generally increases during flash expansion, then decreases during grouping until reaching a constant lowest level during swarming/schooling. This general dynamics is modulated by several parameters: the frequency, direction (vertical vs. horizontal) and target (centroid of the prey aggregation vs. random prey) of predator attacks; the distance at which prey detect predators; the number of prey and predators. Our results suggest that both aerial and underwater predators are more efficient at disturbing fish schools by increasing their attack frequency at such level that the fish cannot return to swarming/schooling. We find that a mix between aerial and underwater predators is more efficient at disturbing a fish school than a single type of attack, suggesting that aerial and underwater foragers may gain mutual benefits in forming foraging groups.     

Sacrificial males: the potential role of copulation and predation in contributing to copepod sex‐skewed ratios

Predation is thought to play a selective role in the emergence of behavioural traits in prey. Differences in behaviour between prey demographics may, therefore, be driven by predation with select components of the population being less vulnerable to predators. While under controlled conditions prey demography has been shown to have consequences for predation success, investigations linking these implications to natural prey population demographics are scarce. Here we assess predator–prey dynamics between notonectid predators (backswimmers) and Lovenula raynerae (Copepoda), key faunal groups in temperate ephemeral pond ecosystems. Using a combination of field and experimental approaches we test for the development and mechanism of predation‐induced sex‐skewed ratios. A natural population of L. raynerae was tracked over time in relation to their predator (notonectid) and prey (Cladocera) numbers. In the laboratory, L. raynerae sex ratios were also assessed over time but in the absence of predation pressure. Predation success and prey performance experiments evaluating differences between L. raynerae male, female, gravid female and copulating pairs exposed to notonectid predation were then examined. Under natural conditions, a female dominated copepod population developed over time and was correlated to predation pressure, while under predator‐free conditions non sex‐skewed prey population demographics persisted. Predator–prey laboratory trials showed no difference in vulnerability and escape performance for male, female and gravid female copepods, but pairs in copula were significantly more vulnerable to predation. This vulnerability was not shared by both sexes, with only female copepods ultimately escaping from successful predation on a mating pair. These results suggest that contact periods during copula may contribute to the development of sex‐skewed copepod ratios over time in ecosystems dominated by hexapod predators. This is discussed within the context of vertebrate and invertebrate predation and how these dissimilar types of predation are likely to have acted as selective pressures for copepod mating systems.     

Is there always an influence of shoal size on predator hunting success?

Theoretical and empirical studies predict that there should be a decrease in hunting success of predators with increasing prey group size. Most of these studies investigated situations in which predator and prey were in full view of each other before, during and after an attack. In this study, single rock bass Ambloplites rupestris were given an opportunity to launch surprise attacks at shoals of creek chub Semotilus atromaculatus that ranged in size from two to 13 fish. There was no significant influence of either shoal size or attack distance on predator success rate and no significant relationship between attack distance and shoal size. Furthermore, it was found that the leading fish of a shoal was attacked significantly more often than fish in other shoal positions, indicating that predation risk was not shared equally among shoal members. Also, leading fish in larger shoals (eight to 13 fish) were not more likely to survive a predator attack than ones in small shoals (two to seven fish).The consequences of these results are discussed in the general context of antipredator benefits of grouping.     

Plasticity of Escape Responses: Prior Predator Experience Enhances Escape Performance in a Coral Reef Fish

Teleost and amphibian prey undertake fast-start escape responses during a predatory attack in an attempt to avoid being captured. Although previously viewed as a reflex reaction controlled by the autonomic nervous system, the escape responses of individuals when repeatedly startled are highly variable in their characteristics, suggesting some behavioural mediation of the response. Previous studies have shown that fishes are able to learn from past experiences, but few studies have assessed how past experience with predators affect the fast-start response. Here we determined whether prior experience with the smell or sight of a predator (the Dottyback, Pseudochromis fuscus) affected the escape response of juveniles of the Spiny Chromis (Acanthochromis polyacanthus). Results show that individuals exposed to any of the predator cues prior to being startled exhibited a stronger escape response (i.e., reduced latency, increased escape distance, mean response speed, maximum response speed and maximum acceleration) when compared with controls. This study demonstrates the plasticity of escape responses and highlights the potential for naïve reef fish to take into account both visual and olfactory threat cues simultaneously to optimise the amplitude of their kinematic responses to perceived risk.     

Plankton reach new heights in effort to avoid predators

The marine environment associated with the air-water interface (neuston) provides an important food source to pelagic organisms where subsurface prey is limited. However, studies on predator-prey interactions within this environment are lacking. Copepods are known to produce strong escape jumps in response to predators, but must contend with a low-Reynolds-number environment where viscous forces limit escape distance. All previous work on copepod interaction with predators has focused on a liquid environment. Here, we describe a novel anti-predator behaviour in two neustonic copepod species, where individuals frequently exit the water surface and travel many times their own body length through air to avoid predators. Using both field recordings with natural predators and high-speed laboratory recordings, we obtain detailed kinematics of this behaviour, and estimate energetic cost associated with this behaviour. We demonstrate that despite losing up to 88 per cent of their initial kinetic energy, copepods that break the water surface travel significantly further than those escaping underwater and successfully exit the perceptive field of the predator. This behaviour provides an effective defence mechanism against subsurface-feeding visual predators and the results provide insight into trophic interactions within the neustonic environment.     

Danger of zooplankton feeding: the fluid signal generated by ambush-feeding copepods

Zooplankton feed in any of three ways: they generate a feeding current while hovering, cruise through the water or are ambush feeders. Each mode generates different hydrodynamic disturbances and hence exposes the grazers differently to mechanosensory predators. Ambush feeders sink slowly and therefore perform occasional upward repositioning jumps. We quantified the fluid disturbance generated by repositioning jumps in a millimetre-sized copepod (Re ∼ 40). The kick of the swimming legs generates a viscous vortex ring in the wake; another ring of similar intensity but opposite rotation is formed around the decelerating copepod. A simple analytical model, that of an impulsive point force, properly describes the observed flow field as a function of the momentum of the copepod, including the translation of the vortex and its spatial extension and temporal decay. We show that the time-averaged fluid signal and the consequent predation risk is much less for an ambush-feeding than a cruising or hovering copepod for small individuals, while the reverse is true for individuals larger than about 1 mm. This makes inefficient ambush feeding feasible in small copepods, and is consistent with the observation that ambush-feeding copepods in the ocean are all small, while larger species invariably use hovering or cruising feeding strategies.     

Predation on the Invasive Copepod,Pseudodiaptomus forbesi,and Native Zooplankton in the Lower Columbia River: An Experimental Approach to Quantify Differences in Prey-Specific Feeding Rates

Invasive planktonic crustaceans have become a prominent feature of aquatic communities worldwide, yet their effects on food webs are not well known. The Asian calanoid copepod, Pseudodiaptomus forbesi, introduced to the Columbia River Estuary approximately 15 years ago, now dominates the late-summer zooplankton community, but its use by native aquatic predators is unknown. We investigated whether three species of planktivorous fishes (chinook salmon, three-spined stickleback, and northern pikeminnow) and one species of mysid exhibited higher feeding rates on native copepods and cladocerans relative to P. forbesi by conducting `single-prey’ feeding experiments and, additionally, examined selectivity for prey types with `two-prey’ feeding experiments. In single-prey experiments individual predator species showed no difference in feeding rates on native cyclopoid copepods (Cyclopidae spp.) relative to invasive P. forbesi, though wild-collected predators exhibited higher feeding rates on cyclopoids when considered in aggregate. In two-prey experiments, chinook salmon and northern pikeminnow both strongly selected native cladocerans (Daphnia retrocurva) over P. forbesi, and moreover, northern pikeminnow selected native Cyclopidae spp. over P. forbesi. On the other hand, in two-prey experiments, chinook salmon, three-spined stickleback and mysids were non- selective with respect to feeding on native cyclopoid copepods versus P. forbesi. Our results indicate that all four native predators in the Columbia River Estuary can consume the invasive copepod, P. forbesi, but that some predators select for native zooplankton over P. forbesi, most likely due to one (or both) of two possible underlying casual mechanisms: 1) differential taxon-specific prey motility and escape responses (calanoids > cyclopoids > daphnids) or 2) the invasive status of the zooplankton prey resulting in naivety, and thus lower feeding rates, of native predators feeding on invasive prey.     

Trophodynamics of the plankton community at Dogger Bank: predatory impact by larval fish

The trophodynamics of a coastal plankton community were studied,focusing on fish larvae and their copepod prey. The major objectiveswere to describe distributional overlap and evaluate the predatoryimpact by larval fish. The study was carried out across DoggerBank in the North Sea, August-September 1991. Sampling transectscrossed tidal fronts off the Bank and plankton at all trophiclevels showed peak abundance within frontal zones. Also Verticallythere was a significant overlap in distributional patterns ofthe plankton. Seven species of fish larvae were abundant, ofthese sprat (Sprattus sprattus) dominated. The abundance ofone group of fish larvae peaked in the shallow water close tothe Bank, whereas other species, including sprat, were foundin deeper water. Prey preference and predation pressure of fishlarvae were assessed using information on prey sizes and growthrates of larvae and the copepod prey. We estimated larval removalof preferred prey sizes to 3–4% day^–1, counterbalancedby a 3–7% day^–1' replenishment from copepod productionand growth. Additional predation pressure on copepods by aninvertebrate predator was estimated to 1–3%day^–1.In conclusion, the dynamics of fish larvae and other zooplankterswere closely linked. At peak abundances of fish larvae (>35mg dry weight m^–2), the accumulated predation on specificsize ranges of copepods, exerted by larvae and other predators,could exceed the ability of copepod replenishment and intra-/interspecificcompetition among predators might take place.     

Detection and capture of inert particles by calanoid copepods: the role of the feeding current

Although there is a scarcity of supporting empirical evidence,it has long been suspected that calanoid copepods use mechanoreceptionto detect the presence and location of potential prey itemsentrained in the feeding current. In this study, we documentthe first observations showing a freely swimming calanoid copepod,Skistodiaptomus oregonensis, attacking prey-sized, non-motile,inert particles entrained in the feeding current before theparticles contact the copepod's sensory appendages. Feedingcurrent geometry, fluid velocities and associated behavioursthat characterize these interactions are described. The resultsof this study show how copepod swimming behaviour, coupled witha low-velocity feeding current, not only increases copepod encounterrates with inert prey by increasing direct contact rates, butalso increases the probability of detecting and capturing remotelylocated prey that have well-developed escape responses. In turbulentregimes, a far-reaching, low-velocity feeding current shouldincrease encounter rates, but only if coupled with behavioursthat quickly minimize separation distances once prey is detected.     

A method for in situ estimation of prey selectivity and predation rate in large plankton, exemplified with the jellyfish Aurelia aurita (L.)

A method to estimate predation rates of large predatory zooplankton, such as jellyfish and ctenophores, is outlined. Large plankton size allows direct visual tracking of the predator during the process of foraging. The presented method is novel in the sense that it measures predation rate of a specific individual plankton predator in situ.After prey has been evacuated from the gut of an individual predator, the predator is incubated in situ, and observed by SCUBA-divers who recapture the individual after a defined time. Given that this incubation time is shorter than prey digestion time, predation rate can be calculated as increase in gut content over time. Clearance rates for different prey can be calculated from predation rates and prey concentrations in the water, allowing accurate estimates of prey selectivity. Thus, the problem of unknown feeding history and feeding environment, which can otherwise be a problem in prey selectivity studies of in situ-captured predators, is circumvented. Benefits and limitations of the method are discussed.The method was applied to adult medusae of the common jellyfish Aurelia aurita. A large variation in number of captured prey was detected both among individual jellyfish and among the various oral arms and gastric pouches within individuals. Clearance rates varied strongly with prey type. The medusae selected large crustacean prey (cladocerans and copepods/copepodites) over echinoderm larvae and copepod nauplii. Prey distribution within the medusae indicates that both tentacles and oral arms were used as prey capturing sites. Food passage time from prey capturing organs to gastric pouches was estimated.     

Threat-Sensitive Responses to Predator Attacks in a Damselfly

The threat sensitivity hypothesis predicts that prey species assess and adjust their behavior flexibly in accordance with the magnitude of the threat imposed by a predator. We tested this hypothesis with regard to escape behavior and thanatosis (feigning of death to escape predation) in larvae of the damselfly Ischnura elegans. We manipulated the perceived predation threat of the larvae by changing three factors: lamellae autotomy (an escape strategy where animals sacrifice a body part when grasped by a predator; lamellae present or absent), kairomone type (odors released by predators; control, dragonfly kairomones or fish kairomones), and population of origin (fishpond or fishless pond). We demonstrated that thanatosis increased survival both when confronted with dragonfly and fish predators. We could show, for the first time, costs of past autotomy to be predator‐dependent: larvae without lamellae suffered higher predation mortality but only in the presence of a dragonfly predator and not in the presence of a fish predator. This is in accordance with the observed reduced escape speed of larvae after autotomy, which may affect escape probability toward dragonfly predators but not to the very fast fish predators. Unexpectedly, kairomone type did not affect the escape response of the larvae. In accordance with the threat sensitivity hypothesis, after an unsuccessful attack, larvae without lamellae had a higher frequency to enter thanatosis than larvae with lamellae and larvae from the fishpond showed longer thanatosis durations than larvae from the fishless pond. Consistent with the hypothesis, the reaction of the larvae to a simulated attack depended jointly on lamellae status and population. In fishless ponds, larvae with lamellae swam away more frequently than larvae without lamellae; in fishponds both groups almost never swam away and relied mostly upon immobility. Given the obvious benefits of adaptively varying escape responses we hypothesize this threat sensitivity to be widespread. Moreover, we argue that former inconsistencies between studies with regard to escape behavior may have been partly because of such adaptive variation.     

Foraging behavior of the predaceous cladoceran, Leptodora kindti, and escape responses of their prey

Using silhouette video photography we have made the first quantitativeobservations of foraging behavior in Leptodora kindti, a predaceouscladoceran (Haplopoda). Leptodora swims with a mean velocityof 13.4?4.0 mm s^–1 and initiates an attack only upon directcontact with potential prey. The attack sequence is as follows:Leptodora swims randomly through the water column with all fivepairs of thorac appendages spread to form a ‘feeding basket’and, seemingly by chance, encounters prey. Shortly after preymake contact with any part of Leptodora's body (usually ventral),the abdomen is rapidly pulled forward, clamping itself underthe feeding basket so that the telson closes it at the posteriorend. The duration of this movement is always the same and weconclude that it is an indiscriminate reflex. If the prey isencountered anywhere but a short distance directly in frontand slightly below the Leptodora, it is not captured. The speedof copepod escape responses effectively allows them to avoidcontact with the predator. Daphnia's escape response, particularlythat of juveniles, is slower and leaves them far more susceptibleto Leptodora predation.     

Movement and direction of movement of a simulated prey affect the success rate in barn owl Tyto alba attack

The present study was aimed at testing a novel idea, that rather than maximizing their distance from a predator during close-distance encounters, prey species are better off moving directly or diagonally toward the predator in order to increase the relative speed and confine the attack to a single available clashing point. We used two tamed barn owls Tyto alba to measure the rate of attack success in relation to the direction of prey movement. A dead mouse or chick was used to simulate the prey, pulled to various directions by means of a transparent string during the owl's attack. Both owls showed a high success rate in catching stationary compared with moving food items (90% and 21%, respectively). Success was higher when the prey moved directly away, rather than towards the owls (50% and 18%, respectively). Strikingly, these owls had 0% success in catching food items that were pulled sideways. This failure to catch prey that move sideways may reflect constraints in postural head movements in aerial raptors that cannot move the eyes but rather move the entire head in tracking prey. So far there is no evidence that defensive behavior in terrestrial prey species takes advantage of the above escape directions to lower rates of predator success. However, birds seem to adjust their defensive tactics in the vertical domain by taking-off at a steep angle, thus moving diagonally toward the direction of an approaching aerial predator. These preliminary findings warrant further studies in barn owls and other predators, in both field and laboratory settings, to uncover fine predator head movements during hunting, the corresponding defensive behavior of the prey, and the adaptive significance of these behaviors.     

Plankton predation rates in turbulence: a study of the limitations imposed on a predator with a non-spherical field of sensory perception

This paper presents an extension to previously published work which studied encounter rates of planktonic predators with restricted perception fields, to examine the related problems of prey capture and predation rates. Small-scale turbulence influences planktonic predation in two ways: the extra energy of the flow enhances the number of encounter events between individual predator and prey meso/micro-zooplankton, but it lowers the capture probability (because the time spent by the predator and prey in close proximity is reduced). Typically, an 'encounter' has usually been defined as an event when a potential prey swims (or is advected) to within a distance R of the predator in any direction. However, there is a considerable body of experimental evidence showing that predators perception fields are far from spherical; often they are wedge shaped (e.g. fish larvae), or strongly aligned with the directions of sensory antennae (e.g. copepods); and this is certain to influence optimal predation strategies. This paper presents a theoretical model which for the first time examines the combined problems of both encounter and capture for a predator with a restricted perception field swimming in a turbulent flow. If such a predator adopts a cruising strategy (continuous swimming, possibly with direction changes) the model predictions suggest that predation rates actually vary little with swimming speed, in contrast to predictions made for spherical perception fields. Consequently, cruising predators are predicted to swim at relatively low speeds whilst foraging. However, application of the model to examine the net energy gain of a typical pause-travel predator (the Atlantic cod larva), does predict the existence of an optimal ratio of the length of pauses to time spent swimming (specifically one pause phase to every two travel phases), in line with experimental observations. Kinematic simulations are presented which support these findings.     

How effective are Mesocyclops aspericornis (Copepoda: Cyclopoida) in controlling mosquito immatures in the environment with an application of phytochemicals?

We investigated the susceptibility of Mesocyclops aspericornis and Aedes aegypti larvae to mosquito larvicidal phytochemicals: piperine and eugenol and the predation efficiency of the adult female M. aspericornis on Ae. aegypti larval instar-I and late II at the sublethal concentration of piperine and eugenol. Both prey and predator were susceptible to both the phytochemicals; however, the lethal concentration of either phytochemicals recorded for Ae. aegypti did not exert mortality on the copepods. The predator’s latent time was significantly longer in the phytochemical medium than control. The encounter frequency was significantly higher in control than in the phytochemical medium. For instar-I larvae, the post-encounter attack probabilities were not affected by the tested phytochemicals. However, for instar-II, the post-encounter attack probability was lower in eugenol treatment than in either the control or piperine. The escape and post-escape survival probabilities of larvae were substantially lower in phytochemical treatments than in the control. However, the overall copepod-imposed mortality did not differ significantly between the control and the phytochemical medium. In conclusion, both the phytochemicals modify the copepod feeding behavior without affecting the copepod incurred total larval mortality. Therefore, application of these phytochemicals along with M. aspericornis for mosquito control is advisable.