Effects of satiation on feeding and swimming behaviour of planktivores

Hunger affects the feeding and swimming behaviour in fish. After 36 h of food deprivation, the feeding and swimming behaviour of Pseudorasbora parva (Cyprinidae) was studied under different prey densities (0.5, 1, 2, 5, 10 and 25 of Daphnia pulex per liter). The initial feeding rates showed marked variations in relation to prey availability. Under high prey densities, the initial feeding rate of fish was higher and subsequently decreased faster, when compared to those feeding under low prey densities. At higher prey densities, two factors were involved: that of higher prey encounter rates and also the attainment of food satiation at a faster rate. Across all prey densities, the feeding rates of fish reached a plateau after satiation. The swimming speed of fish was found to be negatively related to the prey density and a significant change in swimming speed was noted as being directly related to the level of satiation. It was found that the increasing satiation level greatly influenced the handling time and reactive volume of predator, which finally caused reduced feeding rates.     

Functional response of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) to Aphis gossypii Glover (Homoptera: Aphididae) in the Laboratory

Stage specific functional response of Harmonia axyridis (Pallas) to varying densities of Aphis gossypii Glover was examined in a simplified cucumber leaf arena under laboratory conditions. All stages of H. axyridis were isolated individually for 24 h with different prey densities at 25 °C and a photoperiod of 16:8 (L:D) h. The number of prey consumed by the predator was checked at 3, 6, 12, and 24 h. All stages of H. axyridis showed a Type II functional response. Based on the random predator equation, estimated attack rates of H. axyridis at 24 h were 0.0037, 0.0442, 0.3590, 0.3228, and 0.1456, and estimated handling times were 4.1001, 2.4575, 0.7500, 0.2132, and 0.1708 h for the first, second, third, and fourth instars, and female adult, respectively.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

Prey stage preference and functional response of Euseius hibisci to Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae)

The aims of this study were: (a) determine the prey stage preference of female Euseius hibisci (Chant) (Phytoseiidae) at constant densities of different stages of Tetranychus urticae Koch (Tetranychidae), (b) assess the functional response of the predator females to the varying densities of eggs, larvae, or protonymphs of T. urticae, and (c) estimate the functional response of E. hibisci when pollen of Ligustrum ovalifolium was present as well. We conducted experiments on excised pieces of strawberry leaf arenas (Fragaria ananassa) under laboratory conditions of 25 ± 2 °C, 60 ± 5% RH and 12 h photophase. Our results indicated that the predator consumed significantly more prey eggs than other prey stages. Consumption of prey deutonymphs and adults was so low that they were excluded from the non-choice functional response experiments. The functional response on all food items was of type II. The two parameters of the functional response were estimated for each prey type by means of the adjusted non-linear regression model. The highest estimated value a (instantaneous rate of discovery) and the lowest value of T_h (handling time, including digestion) were found for the predator feeding on prey eggs, and a was lowest and T_h highest when fed protonymphs. Using the jack-knife method, the values for the functional response parameters were estimated. The values of a and T_h produced by the model were similar among all prey types except for the eggs, which were different. Using pollen simultaneously with prey larvae decreased the consumption of the latter over the full range of prey densities The suitability of this predator for biological control of T. urticae on strawberry is discussed.     

Effects of spatial scale and foraging efficiency on the predictions made by spatially-explicit models of fish growth rate potential

Synopsis Spatially-explicit modeling of fish growth rate potential is a relatively new approach that uses physical and biological properties of aquatic habitats to map spatial patterns of fish growth rate potential. Recent applications of spatially-explicit models have used an arbitrary spatial scale and have assumed a fixed foraging efficiency. We evaluated the effects of spatial scale, predator foraging efficiency (combined probabilities of prey recognition, attack, capture, and ingestion), and predator spatial distribution on estimates of mean growth rate potential of chinook salmon,Oncorhynchus tshawytscha. We used actual data on prey densities and water temperatures taken from Lake Ontario during the summer, as well as, simulated data assuming binomial distribution of prey. Results show that a predator can compensate for low foraging efficiency by inhabiting the most profitable environments (regions of high growth rate potential). Differences exist in predictions of growth rate potential across spatial scales of observation and a single scale may not be adequate for interpreting model results across seasons. Continued refinements of this modeling approach must focus on the assumptions of stationary distributions of predator and prey populations and predator foraging tactics.     

Geocoris punctipes as a predator of Bemisia tabaci: a laboratory evaluation

Geocoris punctipes (Say), a predaceous lygaeid not previously docmented as a whitefly predator, was tested in the laboratory as a natural enemy of the sweet potato whitefly, Bemisia tabaci (Genn.). Its stalking behavior with whiteflies as prey was similar to that observed with aphids. A previously unobserved behavior was noted that involves the predators using salivary secretions to fasten the wings of prey to various surfaces, allowing labial probing and feeding. Prey consumption as a function of prey number appeared to follow the pattern of the Holling type II functional response. Handling time per prey item ranged from about 180 to 240 seconds. No changes were observed in handling time devoted to earlier versus later catches. Nutritional quality of whiteflies was measured using crude protein, lipids and carbohydrates as criteria. Performance, in terms of predator behavior, total daily handling time, functional response, energy budget and nutritional quality all support the hypothesis that G. punctipes is a promising candidate for biological control of sweet potato whiteflies.     

Time‐to‐kill: measuring attack rates in a heterogenous landscape with multiple prey types

Although spatial heterogeneity of prey and landscapes are known to contribute to variation around predator‐prey functional response models, few studies have quantified these effects. We illustrate a new approach using data from winter movement paths of GPS‐collared wolves in the Rocky Mountains of Canada and time‐to‐event models with competing risks for measuring the effect of prey and landscape characteristics on the time‐to‐kill, which is the reciprocal of attack rate (aN) in a Holling's functional response. We evaluated 13 a priori models representing hypothesized mechanisms influencing attack rates in a heterogeneous landscape with two prey types. Models ranged from variants on Holling's disc equation, including search rate and prey density, to a full model including prey density and patchiness, search rates, satiation, and landscape features, which were measured along the wolf's movement path. Movement rates of wolves while searching explained more of the variation in time‐to‐kill than prey densities. Wolves did not compensate for low prey density by increasing movement rates and there was little evidence that spatial aggregation of prey influenced attack rates in this multi‐prey system. The top model for predicting time‐to‐kill included only search rate and landscape features. Wolves killed prey more quickly in flat terrain, likely due to increased vulnerability from accumulated snow, whereas attack rates were lower when wolves hunted near human‐made features presumably due to human disturbance. Understanding the sources of variation in attack rates provides refinements to functional response models that can lead to more effective predator–prey management in human‐dominated landscapes.     

A functional response model of a predator population foraging in a patchy habitat

1. Functional response models (e.g. Holling's disc equation) that do not take the spatial distributions of prey and predators into account are likely to produce biased estimates of predation rates. 2. To investigate the consequences of ignoring prey distribution and predator aggregation, a general analytical model of a predator population occupying a patchy environment with a single species of prey is developed. 3. The model includes the density and the spatial distribution of the prey population, the aggregative response of the predators and their mutual interference. 4. The model provides explicit solutions to a number of scenarios that can be independently combined: the prey has an even, random or clumped distribution, and the predators show a convex, sigmoid, linear or no aggregative response. 5. The model is parameterized with data from an acarine predator-prey system consisting of Phytoseiulus persimis and Tetranychus urticae inhabiting greenhouse cucumbers. 6. The model fits empirical data quite well and much better than if prey and predators were assumed to be evenly distributed among patches, or if the predators were distributed independently of the prey. 7. The analyses show that if the predators do not show an aggregative response it will always be an advantage to the prey to adopt a patchy distribution. On the other hand, if the predators are capable of responding to the distribution of prey, then it will be an advantage to the prey to be evenly distributed when its density is low and switch to a more patchy distribution when its density increases. The effect of mutual interference is negligible unless predator density is very high. 8. The model shows that prey patchiness and predator aggregation in combination can change the functional response at the population level from type II to type III, indicating that these factors may contribute to stabilization of predator-prey dynamics.     

Biological control ofAphis pomi [Hom.: Aphididae] ofAphidoletes aphidimyza [Dip.: Cecidomyiidae]; a predator-prey model

Predation ofAphis pomi DeGeer [Hom.: Aphididae] byAphidoletes aphidimyza (Rondani) [Dipt.: Cecidomyiidae] was simulated in Michigan apple orchards using a computer and output validated against field data collected from sleeve cages enclosing aphid infested apple terminals. Lower and upper temperature thresholds for development were 2.9 and 35°C for nymphs ofA. pomi with a mean immature developmental period of 162.3 heat units. Median survivorship of adultA. pomi was 364.3 heat units with an average fecundity of 60.7 offspring per female.A. aphidimyza egg and larval lower threshold and developmental periods were 10.5 and 25.5, 8.1°C and 65.5 heat units, respectively. Larval functional response showed Type II behavior with a y-asymptote of 45 aphids killed per predator. Multiple generation model runs performed under different initial predator: prey densities indicated that current critical predator: prey ratios used in the field for control decisions may underestimate predator efficacy.      

Structural sensitivity and resilience in a predator–prey model with density-dependent mortality

Numerous formulations with the same mathematical properties can be relevant to model a biological process. Different formulations can predict different model dynamics like equilibrium vs. oscillations even if they are quantitatively close (structural sensitivity). The question we address in this paper is: does the choice of a formulation affect predictions on the number of stable states? We focus on a predator–prey model with predator competition that exhibits multiple stable states. A bifurcation analysis is realized with respect to prey carrying capacity and species body mass ratio within range of values found in food web models. Bifurcation diagrams built for two type-II functional responses are different in two ways. First, the kind of stable state (equilibrium vs. oscillations) is different for 26.0–49.4% of the parameter values, depending on the parameter space investigated. Using generalized modelling, we highlight the role of functional response slope in this difference. Secondly, the number of stable states is higher with Ivlev's functional response for 0.1–14.3% of the parameter values. These two changes interact to create different model predictions if a parameter value or a state variable is altered. In these two examples of disturbance, Holling's disc equation predicts a higher system resilience. Indeed, Ivlev's functional response predicts that disturbance may trap the system into an alternative stable state that can be escaped from only by a larger alteration (hysteresis phenomena). Two questions arise from this work: (i) how much complex ecological models can be affected by this sensitivity to model formulation? and (ii) how to deal with these uncertainties in model predictions?     

Does pirimicarb affect the voracity of the euriphagous predator, Coccinella undecimpunctata L. (Coleoptera: Coccinellidae)?

Pirimicarb is considered a selective and effective insecticide for the control of aphids and whiteflies. Coccinella undecimpunctata L. is a euriphagous predator autochthonous to the Azores, which feeds preferentially on aphids. The voracity of 4th instars and adults (males and females) of C. undecimpunctata using Aphis fabae Scopoli or Aleyrodes proletella L. as preys was evaluated in laboratory, as well as the impact of pirimicarb on the feeding performance using A. fabae as prey. In the absence of chemical treatment and when the prey was A. proletella, satiation lower limits were estimated on a density of 200 individuals in a 24-h period, for 4th instars, adult females and males of C. undecimpunctata. With A. fabae, satiation was attained when 200, 150 and 100 aphids were provided to 4th instars, adult females and males, respectively. C. undecimpunctata exhibited a type II functional response for both prey species. Fourth instars displayed a lower handling time than the adults; handling times of the adults where higher when A. fabae was the prey and attack rates were sex-dependent, that is, attack rate of females was higher on A. fabae while of males was higher on A. proletella. Voracity of C. undecimpunctata was not significantly affected by pirimicarb; therefore, the use of this insecticide can constitute a complementary component for the integrated management of A. fabae.     

Sources of variation in predation rates at high prey densities: an analytic model and a mite example

Some mathematical techniques for the analysis of satiation-based predation models previously developed by the first author are applied in the present paper to a model by the second author for predation by the predatory miteMetaseiulus occidentalis (Nesbitt). It turns out that for this predator the predation rate should keep increasing at high prey densities as the square root of the prey density,x. This particular shape of the functional response is shown to occur if and only if the upper satiation threshold for prey capture coincides with the maximum gut capacity. The functional response predicted by the model, moreover, is in fair quantitative agreement with predation rates observed by the third author in artificial arenas. A further analysis of the model shows that the variance of the catch should also increase as the square root ofx. This prediction is consistent in a qualitative manner with the continued increase in the variance of the catch. However, quantitatively, the observed variances are even too large to be compatible with any model in which the feeding rate is subject to regulation by a negative feedback. Therefore, the difference between predicted and observed variances is hypothesized to be due to nonhomogeneities in the experimental material. The inferred additional variance component proportional tox accords fairly well with the trend apparent in the data.     

The effect of hunger level on predation dynamics in the spider Nesticodes rufipes: a functional response study

It is well known that a predator has the potential to regulate a prey population only if the predator responds to increases in prey density and inflicts greater mortality rates. Predators may cause such density-dependent mortality depending on the nature of the functional and numerical responses. As spiders are usually faced with a shortage of prey, the killing behavior of the spider Nesticodes rufipes at varying densities of Musca domestica was examined here through laboratory functional response experiments where spiders were deprived of food for 5 (well-fed) or 20 days (hungry). An additional laboratory experiment was also carried out to assess handling time of spiders. The number of prey killed by spiders over 24- and 168-h periods of predator–prey interaction was recorded. Logistic regression analyses revealed the type II functional response for both well-fed and hungry spiders. We found that the lower predation of hungry spiders during the first hours of experimentation was offset later by an increase in predation (explained by estimated handling times), resulting in similarity of functional response curves for well-fed and hungry spiders. It was also observed that the higher number of prey killed by well-fed spiders over a 24-h period of spider–prey interaction probably occurred due to their greater weights than hungry spiders. We concluded that hungry spiders may be more voracious than well-fed spiders only over longer time periods, since hungry spiders may spend more time handling their first prey items than well-fed spiders.     

Experimental duration and predator satiation levels systematically affect functional response parameters

Empirical feeding studies where density‐dependent consumption rates are fitted to functional response models are often used to parameterize the interaction strengths in models of population or food‐web dynamics. However, the relationship between functional response parameter estimates from short‐term feeding studies and real‐world, long‐term, trophic interaction strengths remains largely unexamined. In a critical first step to address this void, we tested for systematic effects of experimental duration and predator satiation on the estimate of functional response parameters, namely attack rate and handling time. Analyzing a large data set covering a wide range of predator taxa and body masses, we show that attack rates decrease with increasing experimental duration, and that handling times of starved predators are consistently shorter than those of satiated predators. Therefore, both the experimental duration and the predator satiation level have a strong and systematic impact on the predictions of population dynamics and food‐web stability. Our study highlights potential pitfalls at the intersection of empirical and theoretical applications of functional responses. We conclude our study with some practical suggestions for how these implications should be addressed in the future to improve predictive abilities and realism in models of predator–prey interactions.     

Tadpole–odonate larvae interactions: influence of body size and diel rhythm

Several studies have shown that prey and predator body size may affect the outcome of predator–prey interactions. However, few studies have taken in account the changes on predator–prey interactions over 24 h. In a tropical freshwater system I evaluated how predator and prey size, and their diel rhythm in activity influenced the interaction between Physalaemus pustulosus tadpoles and dragonfly larvae. Tadpoles of different size classes were exposed to two size classes of the dragonfly larvae Rhionaeschna spec. Feeding trials were conducted during day and night. Tadpole activity showed a diel rhythm and affected size-selective predation of the smallest dragonfly larvae, but not of the larger ones. Predator and prey size had a significant effect on the prey survivorship and prey size had a significant effect on the preference of the predator. The interaction between both factors was significant, indicating that they did not operate independently. I conclude that the predator–prey interactions between odonate larvae and anuran tadpoles were mainly affected by the size of the prey and the predator, and less by the diel activity pattern of the prey.     

Prey: predator ratio dependence in the functional response of a freshwater amphipod

1. First known for their shredding activity, freshwater amphipods also behave as active predators with consequences for prey population regulation and amphipod coexistence in the context of biological invasions. 2. A way to quantify predation is to determine the average consumption rate per predator, also known as its functional response (FR). 3. Although amphipods are gregarious and can display social interactions that can alter per capita consumption rates, previous studies using the FR approach to investigate amphipod predation ignored such potential mutual interference because they did not consider variations in predator density. 4. We investigated the FR of Echinogammarus berilloni feeding on dipteran larvae with joint variations in prey and predator densities. This bivariate experimental design allowed us to estimate interference and to compare the fits of the three main classes of theoretical FR models, in which the predation rate is a function of prey density alone (prey‐dependent models), of both prey and predator densities (predator‐dependent models) or of the prey‐to‐predator ratio (ratio‐dependent models). 5. The Arditi–Ginzburg ratio‐dependent FR model provided the best representation of the FR of E. berilloni, whose predation rate showed a decelerating rise to a horizontal asymptote as prey abundance increased. 6. Ratio dependence means that mutual interference between amphipods leads to prey sharing. Mutual interference is likely to vary between amphipod species, depending on their level of aggressiveness.     

Density-dependent predation ofChaoborus flavicans onDaphnia longispina in a small lake: the effect of prey size

Functional response curves of fourth instar larvae ofChaoborus flavicans preying on two size classes ofDaphnia longispina were examined throughout three summer seasons in a small forest lake. Data for each size class were fitted to Holling's disc equation. The parametersa (attack rate) andTh (handling time) were calculated for each prey size from these curves. Attack rate was greater and handling time was shorter for small (0.77 mm) than for large (1.82 mm)Daphnia. In 1:1 mixture of these prey size classes the predation rates ofChaoborus on smallDaphnia at prey densities above 20 l^–1 were greater than predicted from the single size-class experiments. The observed predation rates on largeDaphnia were lower than predicted at all prey densities. Since both single size-class and two size-class experiments were run during the same period of time the difference in observed and predicted predation rates could not be attributed to seasonal changes in prey preference ofChaoborus larvae. In experiments with a concentrated mixture of lake zooplankton (dominated byD. longispina)Chaoborus preference forDaphnia decreased as prey body size increased. There was no obvious correlation between selectivity coefficients and size-frequency distributions ofDaphnia. When medium-sizedDaphnia were omitted from calculations the preference of small over large prey did not differ significantly from the predictions of the single size-class model.     

Changing handling times during feeding and consequences for prey size selection of 0+ zooplanktivorous fish

The interrelationship of fish size, prey size and handling time within a 15-min feeding period was studied in three size groups of 0 + roach, Rutilus rutilus, and bleak, Alburnus alburnus. Four size classes of cladoceran prey were used to measure changes in feeding rate and handling time from initial rapid feeding to sustained feeding. Observed differences in increase of handling time between prey size classes led to a change in the prey profitability ranking of those size classes within the first 2 min of the experiments. A 1-min feeding period is interpreted as reflecting an intermediate motivational status between extreme hunger and satiation. The use of average handling times for this period revealed a substantial change in prey profitability estimates compared to previous studies which used handling times based on short-term (a few seconds up to 1 min) feeding. It is not the largest prey items a fish can handle and swallow that are most profitable, but prey of intermediate size. By this approach a closer fit between expectations derived from optimal foraging theory and empirical data on prey size selection of 0 + zooplanktivorous fish is qualitatively achieved. Optimal prey size was found to be close the mouth gape width in small fish of 15 mm standard length, decreasing to 50% of mouth gape width in fish of 40 mm standard length.     

Satiation and the functional response: a test of a new model

Abstract. 1. A model of the functional response to prey density is derived to include the reduction in time available for search, T_s , resulting from predator satiation.
2. For larger prey items predator satiation occurs at each prey capture and T_s is reduced by the attack time and digestive pause of a series of attack cycles. For small prey items predator foraging is continuous at low densities with T_s reduced solely by attack time. At higher densities predator satiation occurs after the capture of several small prey items and T_s is reduced by the attack time and digestive pause of a series of foraging cycles.
3. A comparison of the predicted asymptotic level of prey capture using experimentally estimated parameter values, with the maximum consumption of aphids by larval and adult coccinellids provides a test of the satiation model.
4. The limitation of prey capture by predator satiation is discussed with reference to handling time and the success of coccinellids in biological control.