Temperature dependence of the functional response

The Arrhenius equation has emerged as the favoured model for describing the temperature dependence of consumption in predator-prey models. To examine the relevance of this equation, we undertook a meta-analysis of published relationships between functional response parameters and temperature. We show that, when plotted in lin-log space, temperature dependence of both attack rate and maximal ingestion rate exhibits a hump-shaped relationship and not a linear one as predicted by the Arrhenius equation. The relationship remains significantly downward concave even when data from temperatures above the peak of the hump are discarded. Temperature dependence is stronger for attack rate than for maximal ingestion rate, but the thermal optima are not different. We conclude that the use of the Arrhenius equation to describe consumption in predator-prey models requires the assumption that temperatures above thermal optima are unimportant for population and community dynamics, an assumption that is untenable given the available data.     

Pollinator functional response and plant population dynamics: Pollinators as a limiting resource

Summary During recent years, much work has focused on which factors limit the reproductive success in plants. Several studies show a strong influence of either resource limitation, pollen limitation or a combined effect of both. The theoretical arguments for resource limitation are abundant, but there has been very little work done concerning the effect of pollinator availability. In this paper we construct a model to study how the reproductive success in plants is influenced by the foraging behaviour of the pollinators. The pollinator population is assumed to have a constant population density. A functional response function for the pollinators is derived. It is similar to a Holling type II functional response. It is shown that, since the pollinators are regulated by factors not included in the model and their capability to pollinate is limited by the functional response, this is sufficient for regulating the plant population. There also exists a threshold condition for the persistence of the plant population that depended on the search rate of the pollinators and the demographic parameters of the plant population. If this threshold condition is not satisfied the plant population cannot persist and will become extinct. If the condition is satisfied the plant population grows until it is limited at the equilibrium mentioned above.     

Satiation time and predatory behaviour of the dragonfly nymph Mesogomphus lineatus

Summary The dragonfly nymph Mesogomphus lineatus satiated predating 21 larvae of Culex fatigans in 50 minutes; number of attack and predatory efficiency decreased precipitously from 2 attacks/min and 70% during the first 10 minutes of feeding to 0.04 attack/min and 0.01% efficiency respectively during the sixth 10 minutes interval. The nymphs fed after 6, 12, 18, 24, 36 or 42 hours of deprivation consumed 4, 13, 15, 20, 21 or 21 larvae; apparently, the maximum appetite is returned after about 36 hours of deprivation. Satiation time, which was 50 minutes at the density of 15 larvae/aquarium, decreased to 30 minutes in aquaria containing 200 larvae. The nymphs predated increased the number of prey when they were exposed to higher densities of C. fatigans and Anopheles stephansi larvae, and pupa of C. fatigans. They consumed equal weight (but different number) of Culex and Anopheles larvae at all the tested prey densities and selectively selected Culex larva over the pupa or Anopheles larva. Comparative analyses suggest that the dragonfly nymphs deserve serious consideration as larvivorous predators.     

Measuring the functional responses of farmland birds: an example for a declining seed-feeding bunting

1. Many farmland bird species have undergone significant declines. It is important to predict the effect of agricultural change on these birds and their response to conservation measures. This requirement could be met by mechanistic models that predict population size from the optimal foraging behaviour and fates of individuals within populations. A key component of these models is the functional response, the relationship between food and competitor density and feeding rate. 2. This paper describes a method for measuring functional responses of farmland birds, and applies this method to a declining farmland bird, the corn bunting Miliaria calandra L. We derive five alternative models to predict the functional responses of farmland birds and parameterize these for corn bunting. We also assess the minimum sample sizes required to predict accurately the functional response. 3. We show that the functional response of corn bunting can be predicted accurately from a few behavioural parameters (searching rate, handling time, vigilance time) that are straightforward to measure in the field. These parameters can be measured more quickly than the alternative of measuring the functional response directly. 4. While corn bunting violated some of the assumptions of Holling's disk equation (model 1 in our study), it still provided the most accurate fit to the observed feeding rates while remaining the most statistically simple model tested. Our other models may be more applicable to other species, or corn bunting feeding in other locations. 5. Although further tests are required, our study shows how functional responses can be predicted, simplifying the development of mechanistic models of farmland bird populations.     

Dynamics of handling time and functional response by larvae of Chrysomya albiceps (Dipt., Calliphoridae) on different prey species

Abstract:  One way to understand the behavioural patterns exhibited by a predator in response to prey density is to evaluate its functional response. Such evaluation yields information about basic mechanisms of prey–predator dynamics, and is an essential component of prey–predator models. In this paper we analysed experimentally the functional response and the handling time spent by Chrysomya albiceps on different prey species and larval instars of blowflies. The type II functional response was observed when second instar larvae of Chrysomya megacephala and Chrysomya macellaria were consumed. The handling time spent by the predator was significantly different between instars and species. The implications of the functional response and handling time for the interaction dynamics of Brazilian Chrysomyinae species are discussed.     

Harvest rates and foraging strategies in Negev Desert gerbils

We examined the foraging strategy and quantified the foragingtraits of two nocturnal rodent species, Allenby's gerbil (Gerbillusallenbyi) and the greater Egyptian sand gerbil (Gerbillus pyramidum).In the laboratory, both species used two distinct foragingstrategies: either they immediately consumed seeds found ina patch (seed tray); or they collected and delivered the seedsto their nest box for later consumption. Moreover, we founda transition in foraging strategy among individual G. allenbyi under laboratory conditions; they all began by consuming theseeds on the tray and, after 7 days on average, switched tothe collecting strategy. By contrast, in the field both speciesused only one foraging strategy; they collected and deliveredthe seeds to their burrow or to surface caches for later consumption.Furthermore, G. allenbyi and G. pyramidum collected seeds atsignificantly higher rates in the field than in the laboratorybecause the seed encounter rates for both species were higherin the field. This suggests that in natural conditions, probablyinvolving predation risk and competitive pressure, gerbilsmust respond in two ways: (1) they must choose a foraging strategythat reduces predation risk by minimizing time spent feedingoutside their burrows; and (2) they must forage more efficiently.In the field, seed handling time of the larger species, G. pyramidum, was shorter than that of the smaller one, G. allenbyi.This difference may give G. pyramidum an advantage when resourcelevels are high and when most of a forager's time is spent handling seeds rather than searching for more seeds. Additionally,our field study showed that the seed encounter rate of G. allenbyiwas higher than that of G. pyramidum. This difference may giveG. allenbyi an advantage when resource levels are low and whensearching occupies most of the forager's time. The differentadvantages that each species has over the other, under differentconditions, may well be factors promoting their coexistenceover a wide range of resource densities.     

Effect of treating eggs of cotton bollworm with Bacillus thuringiensis Berliner on functional response of Trichogramma brassicae Bezdenko

Cotton bollworm, Helicoverpa armigera (Hübner) is a cosmopolite insect pest of a wide spectrum of crops such as cotton, maize, tomato, soybean, etc. Egg parasitoids mainly Trichogramma brassicae Bezdenko and Bacillus thuringiensis Berliner (Bt) are biological control agents, that are used as components of sustainable and environmentally compatible IPM systems. Although Bt does not come in direct contact with egg parasitoids, it may persist within the host’s body and affect the quality of the host’s eggs via biochemical changes in their mother and possibly behaviour and potency of the parasitoids. In this study, the functional response of T. brassicae to different densities of H. armigera eggs was investigated in two sets of experiments at 26?±?1?°C, 65?±?5% RH, and 16: 8?h photoperiod. The first group was a control and the second one were eggs laid by hosts treated as 3rd instar larvae with LC₂₀ of Bt (determined as 9.8?×?10^5?IU/l of artificial diet based on a preliminary bioassay). A type III functional response was observed in both treatments with a direct density dependent mortality up to eight host eggs and an inverse one upward. Both handling time and searching efficiency were affected by Bt treatment as the handling time was increased by a factor of 1.5 and the searching efficiency was decreased by a factor of 0.6. The searching efficiencies were 0.0310?±?0.003 and 0.0182?±?0.005?h^?1, and handling times were 1.134?±?0.042 and 1.672?±?0.082?h in control and Bt treatment respectively.     

Predatory responses of selected lines of developmental variants of ladybird,Propylea dissecta (Coleoptera: Coccinellidae) in relation to increasing prey and predator densities

Individuals of the same species, population and generation frequently exhibit sub-maximal and significant genetic and phenotypic variation in their rate of development, showing slow and fast developers. Fast developers commonly have higher foraging and predation rates than slow developers. The consequence of such differences and foraging for the efficacy of biocontrol species remains under-explored. Slow and fast developers from a population of the ladybird, Propylea dissecta were separated and selected experimentally for F15 generations, and the predatory response of fourth instar larvae of control and experimentally selected slow and fast developers was then assessed at differing levels of prey (pea aphid, Acyrthosiphon pisum) and conspecific predator abundance. All individuals, whether slow or fast developers, showed a Type-II functional response, decrease in proportion of prey consumed with increasing prey biomass and an increase in proportion of prey consumed with increasing predator density. The proportion of prey consumed was highest in experimental fast developers and lowest in experimental slow developers. Attack rate was highest and handling time longest in slow developers of control/experimental groups. Mutual interference was least while area of discovery was highest in experimental fast developers. Thus, selection of fast developers for F15 generations led to higher functional responses, slower attack rates and faster prey consumption. This lower mutual interference and high searching efficiency indicates that they can be experimentally selected and used for better control of the pea aphids. This study is the first attempt to evaluate predatory responses of selected lines of an aphidophagous ladybird.     

The effect of prey species and environmental complexity on the functional response of Franklinothrips orizabensis: a test of the fractal foraging model

Abstract. 1. The hypothesis proposed by the fractal foraging model, that prey procurement by predators declines as habitat complexity increases, was tested. To evaluate this hypothesis, the effect of two prey species, second-instar larvae of Scirtothrips perseae Nakahara and Heliothrips haemorrhoidalis (Bouché) (both Thysanoptera: Thripidae), and environmental complexity on the functional response of the predatory thrips Franklinothrips orizabensis Johansen (Thysanoptera: Aeolothripidae) was examined.
2. The Koch curve, a well-studied fractal, was used to generate the shapes of experimental arenas to test the effect of environmental complexity on the functional response of F. orizabensis. Complexity was defined in terms of the number of acute vertices in which prey thrips could seek refuge and the length of the perimeter that had to be searched by the predator. Four shapes were tested: a circle (zero acute vertices, circumference = 186.61 mm), equilateral triangle (three acute vertices, perimeter = 240 mm), star (six acute vertices, perimeter = 277.13 mm), and snowflake (18 acute vertices, perimeter = 333.65 mm). All shapes were scaled so that the area of each arena was 2771 mm².
3.  Franklinothrips orizabensis exhibited a Type II functional response for both prey species and all four experimental arenas tested. Significantly fewer S. perseae larvae were killed in the most complex arena (i.e. the snowflake) when prey densities exceeded 16. For H. haemorrhoidalis , significant differences in mortality were observed only when prey densities equalled four. These results demonstrated that the fractal foraging model was supported when certain prey densities coincided with particular levels of environmental complexity.
4. Subtle changes in environmental complexity and different prey species of varying density can affect prey–predator interactions significantly.     

Towards a mechanistic understanding of individual-level functional responses: Invasive crayfish as model organisms

1. In novel communities, a rising number of new and emerging invasive species interact with resident species, some of which are non-native themselves. We implemented an innovative trophic interaction framework for novel communities and quantified the interaction strength and impact potential of a truly novel species (marbled crayfish Procambarus virginalis) with a resident non-native counterpart (spiny-cheek crayfish Faxonius limosus). As prey, we used Dreissena mussels, which are non-native as well and now hold a key position in many European and North American aquatic ecosystems.
2. For both crayfish species, we predicted functional responses based on a mechanistic model that we parameterised with a set of experimental observations of foraging behaviour and satiation. We compared these predicted functional responses to empirically observed responses. In addition, we incorporated behavioural traits such as aggression, activity, and boldness in the comparisons between the species and individuals to determine their influence on functional responses. We tested individuals from aquarium stocks as well as naturalised individuals from invaded water bodies.
3. Altogether, we performed 1,095 experiments with 26 individual crayfish. We found that per capita predation of spiny-cheek crayfish exceeded that of marbled crayfish from aquaria and naturalised individuals. Functional responses differed between species and were mostly higher for spiny-cheek crayfish males. Marbled crayfish, however, were more voracious and reached satiation more slowly. Consumption rates correlated with aggression for marbled crayfish and with an aggressive threat response for spiny-cheek crayfish.
4. We conclude that spiny-cheek crayfish can reach higher short-term consumption rates than marbled crayfish, but both species probably do not substantially affect Dreissena mussel populations in the field. For marbled crayfish, high long-term consumption, interspecific aggression, and reproduction rates can promote their establishment and spread. Risk assessments of these invaders should be improved by considering numerical responses, and different prey organisms and predators.

     

Intake rates and the functional response in shorebirds (Charadriiformes) eating macro-invertebrates

As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II ('disc equation') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching.A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m-2). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote.A multivariate analysis of 468 'spot' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6%, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases. We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested. As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.     

Assessing superparasitism with a model combining the functional response and the egg distribution of parasitoids

A model for superparasitism in insect parasitoids is developed. This model combines the study of superparasitism in terms of distribution of eggs among hosts (for a given number of hosts) and in terms of functional response (number of hosts attacked for a varying number of hosts available). Thus, it gives a synthetic treatment of problems that had been previously handled with separate models (e.g., Bakkeret al. (1972) on one hand and Arditi (1983) on the other hand). The combined model involves several parameters, among which important ones are the propensity to superparasitise, δ, and the average handling times spent on healthy and parasitised hosts, T_h and T_p. Special cases are those of an indiscriminate parasitoid (δ=1 and T_p=T_h) and of a “predator-like” parasitoid (δ=0 and T_p=0). In this paper, the emphasis is put on the problems related with model identification and parameter estimation from experimental data. According to the data available, three situations are considered: egg distribution alone, functional response alone, and both combined. The main conclusions are the following. (i) Egg distributions are described correctly when the parasitoid/host ratio is not too high. When the situation is very strained, i.e., when a small number of hosts are available per parasitoid, superparasitism occurs more frequently than predicted by the model. (ii) Functional response data are usually not precise enough to estimate all model parameters, particularly T_p. That is, it will usually not be possible to assess the discrimination capacity of a given species on the basis of a functional response curve only. (iii) If both a functional response and the corresponding egg distributions are available, it is better to fit the egg distribution model first and, depending on the estimated value of δ, to fit thereafter the appropriate functional response model.     

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.