Effect of tomato leaf hairiness on functional and numerical response of Neoseiulus californicus (Acari: Phytoseiidae)

The objective of this study was to determine whether differences in hairiness of tomato plants affect the functional and numerical response of the predator Neoseiulus californicus McGregor attacking the two-spotted spider mite, Tetranychus urticae Koch. Two tomato hybrids with different density of glandular hairs were used. The functional response was measured by offering eggs and adults of T urticae at densities ranging from 4 to 64 items per tomato leaflet (surface ca. 6.3 cm2); eggs were offered to predator protonymphs and deutonymphs, adult spider mites to adult predators. The number of spider mites eaten as a function of initial density was fitted to the disc equation. Predator densities were regressed against initial prey densities to analyze the numerical response. The number of eggs and adults of T. urticae eaten by N. californicus was extremely low in both hybrids. The nymphal stage of N. californicus and prey density had a significant effect on the number of T urticae eggs eaten by the predator, while hybrid had no effect. The functional response fitted reasonably well to the Holling model. The handling time (Th) and the attack rate (a) were very similar among the two hybrids. The numerical response indicated that the absolute density of predators increased with changes in spider mite densities but the relative predator/prey density decreased in both hybrids. Tomato hairiness prevented N. californicus from exhibiting a strong numerical response and the predator functional response was much lower than observed in other host plants and other phytoseiids. This result shows the need to consider plant attributes as an essential and interactive component of biological control practices.     

Ontogenetic shifts in intraguild predation on thrips by phytoseiid mites: the relevance of body size and diet specialization

In greenhouse agroecosystems, a guild of spider mite predators may consist of the oligophagous predatory mite Phytoseiulus persimilis Athias-Henriot, the polyphagous predatory mite Neoseiulus californicus McGregor (both Acari: Phytoseiidae) and the primarily herbivorous but facultatively predatory western flower thrips Frankliniella occidentalis Pergande (Thysanoptera: Thripidae). Diet-specialization and the predator body size relative to prey are crucial factors in predation on F. occidentalis by P. persimilis and N. californicus. Here, it was tested whether the relevance of these factors changes during predator ontogeny. First, the predator (protonymphs and adult females of P. persimilis and N. californicus): prey (F. occidentalis first instars) body size ratios were measured. Second, the aggressiveness of P. persimilis and N. californicus towards F. occidentalis was assessed. Third, survival, development and oviposition of P. persimilis and N. californicus with F. occidentalis prey was determined. The body size ranking was P. persimilis females > N. californicus females > P. persimilis protonymphs > N. californicus protonymphs. Neoseiulus californicus females were the most aggressive predators, followed by highly aggressive N. californicus protonymphs and moderately aggressive P. persimilis protonymphs. Phytoseiulus persimilis females did not attack thrips. Frankliniella occidentalis larvae are an alternative prey for juvenile N. californicus and P. persimilis, enabling them to reach adulthood. Females of N. californicus but not P. persimilis sustained egg production with thrips prey. Within the guild studied here, N. californicus females are the most harmful predators for F. occidentalis larvae, followed by N. californicus and P. persimilis juveniles. Phytoseiulus persimilis females are harmless to F. occidentalis.     

Effect of Prey and Predator Density on Predation of Rice Leaffolder Eggs by the Cricket Metioche vittaticollis

Cage experiments were conducted to quantify the predation rate of the cricket Metioche vittaticollis (Sta ˚l) on the eggs of rice leaffolder Marasmia patnalis Bradley. Egg predation by adult females was measured in response to changes in egg density, predator density and leaf area per cage. The number of eggs consumed per predator increased with egg density, without reaching a plateau. The predation rate decreased with increasing leaf area. The functional response could be adequately described with a linear Type I model, with the effect of leaf area included. This type of response to leaffolder egg density means that predation was not limited by prey handling time or satiation, but by the search rate. The search rate is here interpreted as the leaf area effectively searched for leaffolder eggs by a single predator in one day. Estimated search rates averaged 0.13 m 2 day -1 for M. vittaticollis females. The search rate of the predators increased with prey density, but a model describing a density dependent search rate explained only 3% of the total variation in search rate. Increasing predator density per cage led to a decrease in the per capita egg predation rate when predator density was more than two per m 2 leaf area. Interference might thus reduce the potential to enhance leaffolder egg predation by conservation or augmentation of predatory cricket populations.     

Foraging behavior of the coccinellid Nephus includens (Coleoptera: Coccinellidae) in response to Aphis gossypii (Hemiptera: Aphididae) with particular emphasis on larval parasitism

This study assessed the effect of parasitism of Nephus includens (Col.: Coccinellidae) larvae by Homalotylus flaminius (Hym.: Encyrtidae) on the predation rates of the predator on the cotton aphid, Aphis gossypii Glover (Hem.: Aphididae) by deriving functional responses for second- and fourth-instar predators at prey densities ranging from 10 to 80 aphids per arena. The relationship between the functional and numerical responses of adult females of N. includens also was determined for prey densities ranging from 10 to 140 aphids per arena. Predation rates of unparasitized and parasitized second-instar N. includens were both fit by a type II functional response model with parameters as follows: unparasitized (a = 0.0768 hours(-1) and T(h) = 0.975 h) and parasitized (a = 0.0787 hours(-1) and T(h) = 0.8823 hours). Predation rates of unparasitized and parasitized fourth-instar N. includens were fit by type III and II models, respectively, with the following parameters: unparasitized (b = 0.1702 hours(-1) and T(h) = 0.2369 hours) and parasitized (a = 0.038 h(-1) and T(h) = 0.539 h). The unparasitized fourth-instar was the most voracious stage, having the highest attack rate and lowest handling time. Considering these attributes, it would seem to be the most effective stage of this predator against A. gossypii. Adult female lady beetles (N. includens) showed a type III functional response and their numerical response increase to prey density was curvilinearly related to prey density, with the highest number of eggs being produced at highest prey densities. The maximum saturation level for both predation and egg production for adult females occurred at a prey density of 120 aphids. Thus, a ratio 1:120 (predator:prey) should be used when releasing this species for augmentative biological control. Release of either fourth-instar or adult stage N. includens should be minimized the potentially negative effect of parasitism by H. flaminius on early developmental stages, and hence increase its efficiency in biocontrol programs.     

Diet‐dependent intraguild predation between the predatory mites Neoseiulus californicus and Neoseiulus cucumeris

Based on the hypothesis that matching diets of intraguild (IG) predator and prey indicate strong food competition and thus intensify intraguild predation (IGP) as compared to non‐matching diets, we scrutinized diet‐dependent mutual IGP between the predatory mites Neoseiulus cucumeris and N. californicus. Both are natural enemies of herbivorous mites and insects and used in biological control of spider mites and thrips in various agricultural crops. Both are generalist predators that may also feed on plant‐derived substances such as pollen. Irrespective of diet (pollen or spider mites), N. cucumeris females had higher predation and oviposition rates and shorter attack latencies on IG prey than N. californicus. Predation rates on larvae were unaffected by diet but larvae from pollen‐fed mothers were a more profitable prey than those from spider‐mite fed mothers resulting in higher oviposition rates of IG predator females. Pollen‐fed protonymphs were earlier attacked by IG predator females than spider‐mite fed protonymphs. Spider mite‐fed N. californicus females attacked protonymphs earlier than did pollen‐fed N. californicus females. Overall, our study suggests that predator and prey diet may exert subtle influences on mutual IGP between bio‐control agents. Matching diets did not intensify IGP between N. californicus and N. cucumeris but predator and prey diets proximately influenced IGP through changes in behaviour and/or stoichiometry.     

Euseiusfinlandicus (Acari: Phytoseiidae) as a potential biocontrol agent against Tetranychus urticae (Acari: Tetranychidae): life history and feeding habits on three different types of food

Life history and reproductive parameters of the generalist predatory mite Euseius (Amblyseius) finlandicus (Oudemans) were studied in the laboratory at 25 +/- 1 degrees C, with a 16L:8D photoperiod and 60 +/- 15% RH, to investigate its response to different food sources: an eriophyid mite Aceria sp., tulip pollen Tulipa gesnerana L., and two-spotted spider mite Tetranychus urticae Koch. Total developmental time of the immature stages was the shortest on eriophyid mites, followed by pollen, and then spider mites. Fecundity was highest on pollen (43.69 eggs; 1.63 eggs/female/day), then eriophyid mites (39.73 eggs; 1.37 eggs/female/day) and lowest on spider mites (18.16 eggs; 0.80 eggs/female/day). Intrinsic rate of increase (Rm), net reproductive rate (Ro) and finite rate of increase (lambda) followed the same pattern [pollen (0.168, 27.96 and 1.183, respectively), eriophyid mites (0.153, 20.81 and 1.167), spider mites (0.110, 9.44 and 1.119)]. Mean generation time (days) was the shortest on pollen (19.90), followed by eriophyid mites (20.02), and then spider mites (20.59). Average spider mite larvae consumed by E. finlandicus during immature stages were 9.18 for males and 11.85 for females. Adult E. finlandicus females consumed an average of 166.38 spider mite protonymphs during adult stage compared to an average of 66.55 by males. The number of prey protonymphs consumed per day by females was highest in the oviposition period, lower in the pre-oviposition period and the lowest in the post-oviposition period. The eriophyid mite as a prey recorded the shortest developmental time, while pollen as food recorded the highest oviposition rate in E. finlandicus. The potential of this predator as a biocontrol agent against T. urticae is discussed.     

Prey stage preference and functional response of the predatory mite Galendromus flumenis to Oligonychus pratensis

The Banks grass mite, Oligonychus pratensis (Banks) (Acari: Tetranychidae), is a serious pest in dates (Phoenix dactylifera L.) in the New World. Currently O. pratensis is managed using the miticide, Savey, and alternative strategies are necessary to remove pressure from a single control method due to the risk of resistance evolution. For this purpose, studies are underway to develop biological control strategies using the predatory mite, Galendromus flumenis (Chant) (Acari: Phytoseiidae). The current study determined the consumption rate of G. flumenis at constant densities of O. pratensis eggs, larvae, protonymphs and deutonymphs, and defined the functional response of predator females. The predator consumed significantly more eggs than other prey stages, and displayed a type II functional response on all prey stages. The highest attack rate and shortest handling time were obtained for predators feeding on prey larvae and eggs, respectively. The proportions of prey consumed by G. flumenis were higher at lower densities for all stages of Banks grass mite, implying that G. flumenis should be more effective at suppressing Banks grass mite populations at lower densities. Therefore, in an augmentative release program, G. flumenis would need to be released early in the infestation.     

Behavioral responses to prey density by three acarine predator species with different degrees of polyphagy

Behavioral responses by three acarine predators, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to different egg and webbing densities of the spider mite Tetranychus urticae (Acari: Tetranychidae) on rose leaflets were studied in the laboratory. Prey patches were delineated by T. urticae webbing and associated kairomones, which elicit turning back responses in predators near the patch edge. Only the presence of webbing affected predator behavior; increased webbing density did not increase patch time. Patch time increased with increased T. urticae egg density in the oligophagous P. persimilis, but was density independent in the polyphagous species T. occidentalis and A. andersoni. Patch time in all three species was more strongly correlated with the number of prey encounters and attacks than with the actual prey number present in the patch. Patch time was determined by (a) the turning back response near the patch edge; this response decayed through time and eventually led to the abandonment of the patch, and (b) encounters with, and attacks upon, prey eggs; these prolonged patch time by both an increment of time spent in handling or rejecting prey and an increment of time spent searching between two successive prey encounters or attacks. Although searching efficiency was independent of prey density in all three species, the predation rate by P. persimilis decreased with prey density because its searching activity (i.e. proportion of total patch time spent in searching) decreased with prey density. Predation rates by T. occidentalis and A. andersoni decreased with prey density because their searching activity and success ratio both decreased with prey density. The data were tested against models of predator foraging responses to prey density. The effects of the degree of polyphagy on predator foraging behavior were also discussed.     

Resource-mediated impact of spider predation risk on performance in the grasshopper Ageneotettix deorum (Orthoptera: Acrididae)

In response to increased exposure to predators when searching for food, many prey increase the frequency of antipredator behaviors, potentially reducing foraging rate and food intake. Such direct, nonlethal interactions between predators and prey resulting in reduced food intake can indirectly influence lifecycle development through effects on growth, developmental rate, and survival. We investigated the general hypothesis that individual performance of a herbivorous insect can be negatively affected when exposed to nonlethal predation risk, and that the response can be mediated by food quality. This hypothesis was tested using the common rangeland grasshopper Ageneotettix deorum with and without exposure to common wolf spider predators (Lycosidae, Schizocosa spp.) on both untreated natural and fertilized vegetation. All spiders were rendered temporarily incapable of direct feeding by restricting function of the chelicerae with beeswax. Detectable responses by grasshoppers to spiders indicate indirect consequences for lifecycle development. Grasshopper performance was measured as hind femur growth, duration of nymphal lifecycle stages, and survivorship in a caged field experiment conducted over 2 years. Grasshoppers developed faster and grew 3–5% larger when allowed to forage on fertilized vegetation in the absence of risk from a spider predator. Failure-time analysis illustrated enhanced survival probability in response to elevated food quality and the negative effects of grasshopper susceptibility to nonlethal predation risk. Performance on food of relatively low, ambient quality with no predation risk equaled that of grasshoppers caged with high quality vegetation in the presence of a modified spider. Increased resource quality can clearly moderate the negative life history responses caused by the behavioral modification of grasshoppers when exposed to spider predation risk, a compensatory response.     

Biology of three species of Agistemus (Acari: Stigmaeidae): life table parameters using eggs of Panonychus citri or pollen of Malephora crocea as food

The biology and life table parameters of Agistemus industani Gonzalez, A. cyprius Gonzalez, and A. floridanus Gonzalez (Acari: Stigmaeidae) were studied under laboratory conditions using two food sources: Panonychus citri (McGregor) eggs or ice plant, Malephora crocea (Jacquin) Schwantes pollen at 25 degrees C. The larval, protonymph, deutonymph, and adult stages of A. industani fed on citrus red mite eggs. All active stages of A. industani, except the larva, fed on all P. citri stages and the larval stage could not feed on P. citri adults. All immature stages of A. industani fed on M. crocea pollen. Agistemus cyprius larvae fed on P. citri eggs and larvae or ice plant pollen. The nymphal stages fed on P. citri eggs, larvae, and protonymphs but not deutonymphs or adults while A. cyprius deutonymphs and adults fed on all P. citri stages. Adult and nymphal stages of A. cyprius fed on ice plant pollen and successfully completed their development while A. floridanus did not. Agistemus floridanus larvae fed only on P. citri eggs, while the other stages fed on P. citri eggs, larvae, and protonymphs. The developmental times from egg to adult for A. industani and A. cyprius when fed M. crocea pollen were 11.3 and 13.4 days, respectively. Agistemus floridanus was unable to complete its life cycle on a diet of only M. crocea pollen. Agistemus industani, A. cyprius, and A. floridanus completed development from egg to adult in 11.7, 13.8, and 10.8 days, respectively, when fed P. citri eggs. The intrinsic rate of increase (r(m)) values for A. cyprius and A. industani were 0.0311 and 0.1201 per day on the pollen diet. The net reproductive rate (Ro) was 3.58 for A. cyprius and 10.07 for A. industani with generation times (T) of 45.2 and 35.1 days, respectively, on the ice plant pollen diet. The r(m) values for A. cyprius, A. floridanus, and A. industani on the P. citri egg only diet were: 0.0562, 0.1001, and 0.1031 per day, respectively. The Ro values for each species fed P. citri eggs only were: 6.36, 7.90, and 18.70 for A. cyprius, A. floridanus, and A. industani and the generation times (T) for each of the three species were: 35.2, 29.9 and 37.8 days, respectively.     

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.     

Comparative toxic and sublethal effects of fluvalinate on two-spotted spider mite and European red mite

Four fluvalinate formulations differed in their residual toxicity to female two-spotted spider mite (TSM), Tetranychus urticae adults; the emulsifiable concentrate (EC) was the most toxic. In contrast, there was little difference in toxicity between the formulations with the European red mite (ERM) Panonychus ulmi with the exception of the EC formulation which was the least toxic. Fluvalinate 2F caused minimal (<10%) TSM and ERM egg mortality. Fluvalinate 2F was more toxic and caused greater larval dispersal for the TSM compared to the ERM at the field concentration and below. The toxicity of fluvalinate 2F to TSM and ERM protonymphs, deutonymphs and adults was low, approximately <20% at field concentration. Dispersal was the main response to fluvalinate and this was positively correlated with increasing concentration. The combined mortality and dispersal LC₅₀ was five times lower for ERM protonymphs and adults, but 11 times higher for ERM deutonymphs compared to equivalent TSM life stages. Fluvalinate 2F reduced TSM development from the protonymph and deutonymph stages to a greater extent compared to the ERM. The mortality response to fluvalinate 2F was unaffected by host type (peach or apple) for the TSM whereas ERM mortality was higher on apple compared to peach. TSM dispersal was higher from apple compared to peach whereas ERM dispersal was similar on both host types. Oviposition by both mite species was lower on apple than peach leaves. A 1 h exposure to fluvalinate 2F reduced ERM oviposition for 12 days.     

Positive indirect effect of aquatic insects on terrestrial prey is not offset by increased predator density

1. Changes in one prey species' density can indirectly affect the abundance of another prey species if a shared predator eats both species. Sometimes, indirect effects occur when prey straddle habitats, including when riparian predator populations grow in response to emergent aquatic insects and increase predation on terrestrial prey. However, predators may largely switch to aquatic insects or become satiated, reducing predation on terrestrial prey. 2. To determine the net indirect effect of aquatic insects on terrestrial arthropods via generalist spider predators, a field experiment was conducted mimicking midge influx and a wolf spider numerical response inside enclosures near an Icelandic lake. Lab mesocosms were also used to assess per capita rates of spider predation u nder differing levels of midge abundance. 3. Midges always decreased sentinel prey predation, but this effect increased with predator density. When midges were absent, predation increased 30% at a high spider density, but predation was equal between spider treatments when midges were present. In situ arthropods showed no effect of midge or spider treatments, although non‐significant abundance patterns were observed congruent with sentinel prey results. 4. In lab mesocosms, prey survivorship increased ≥50% where midges were present and rapidly saturated; the addition of 5, 20, 50, and 100 midges equivalently reduced spider predation, supporting predator distraction rather than satiation as the root cause. 5. The present results demonstrate a strong positive indirect effect of midges and broadly support the concept that predator responses to alternative prey are a major influence on the magnitude and direction of predator‐mediated indirect effects.     

A comparative life history study of immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae) with a review of larval feeding patterns in the family

Survival, developmental time, activity, feeding rates, and other biological aspects of immatures of Amblyseius fallacis, Amblyseius andersoni, Typhlodromus occidentalis and Typhlodromus pyri were examined in the laboratory in small arenas (2×2 cm) with different egg densities (0, 5, 10, 20 per 12 h) of the twospotted spider mite, Tetranychus urticae (Koch), at 25±1°C, ≈80% RH, and 16L: 8D photoperiod. Egg survival was high (86–100%) in all four species. Larval survival was similarly high except for T. occidentalis which all died in the absence of food. Survival rates of protonymphs and deutonymphs were also high except that up to 50% of A. andersoni died at 5 prey eggs per 1/2 day. Developmental time did not vary significantly with prey density and was similar for males and females in the oligophagous predators (A. fallacis and T. occidentalis), but was longer at lower prey densities and in females than males in the polyphagous predators (A. andersoni and T. pyri). In general, the time allocated to three active instars (=stases) decreased in the order: A. andersoni (81%), T. pyri (78%), A. fallacis (69%), and T. occidentalis (64%). The polyphagous predator species had a shorter larval stage and much longer deutonymphal stage than the oligophagous species. The proportion of time allocated to the protonymphal stage was the least variable among the four species. The interspecific differences in walking activities also appeared greater in larval and deutonymphal stages than in the protonymphal stage. The larvae of the two oligophagous predators (A. fallacis and T. occidentalis, walking activity averaging 36–49%) were more active than the two polyphagous predators (A. andersoni and T. pyri), which spent 80% or more time resting. In deutonymphs, walking activity increased in the order: T. occidentalis (14%), A. andersoni (27%), A. fallacis (43%) and T. pyri (59%). Larvae were more active during the first half of their life than the latter half. In general, most life history traits of immature A. andersoni, T. pyri, A. fallacis, and T. occidentalis are not associated with their phylogenetic relatedness or size, but with the feeding specialization of the predator species. Larval feeding patterns in Phytoseiidae are reviewed and a hypothesis about the evolution of larval feeding behavior in Phytoseiidae is proposed.     

Host plant effects on the functional response of Stethorus gilvifrons to strawberry spider mites

The functional response of adult females of the coccinellid beetle Stethorus gilvifrons Mulsant to juveniles of strawberry spider mite, Tetranychus turkestani Ugarov & Nikolski was determined on cowpea, castor bean and cucumber leaves in the laboratory at 25°C and a 14 h L: 10 h D photoperiod. Beetles were isolated singly for 24 h in 9-cm Petri dishes with either 2, 4, 8, 16, 32, 64, or 128 nymphal stages of T. turkestani. Results showed a typical type II response on all plants tested, with up to 110.7, 100.8, and 53.0 prey attacked when 128 nymphal stages were provided on cowpea, castor bean, and cucumber leaves, respectively. Based on the Rogers random attack equation, the highest estimated attack rate and the lowest handling time were obtained on cowpea. It was therefore concluded that the host plant species can affect the predation rate and functional response components of S. gilvifrons, a specific and effective predator of spider mites.     

Interaction between life stages in a phytoseiid predator: western flower thrips prey killed by adults as food for protonymphs of Amblyseius cucumeris

Alternative feeding strategies are important in determining the lifestyle of polyphagous spider mite prodators, and could play a key role in their use for biocontrol of prey such as thrips. The small size of Amblyseius cucumeris relative to western flower thrips, Frankliniella occidentalis, limits its survival and development when this prey is the only food available. We show that when A. cucumeris nymphs were reared either alone or with a gravid female on live larvae of F. occidentalis as the only source of food, survival was increased and development was accelerated by the presence of the adult. Similar performance by predator nymphs reared alone on freshly killed thrips larvae indicated that those nymphs reared on live prey with an adult were benefiting from feeding on prey killed by the adult. Variation of the period when an adult female was present with the nymph showed that food provided as a result of the adult's preying activities was beneficial until approximately one third through nymphal development, after which protonymphs became independent predators, with good survival and rapid development when provided only with live F. occidentalis larvae. The results are discussed in relation to adult dispersal in specialist as opposed to generalist phytoseiids, and its potential manipulation in using A. cucumeris for thrips biocontrol.     

Do functional responses of predatory arthropods reach a plateau? A case study of Orius insidiosus with western flower thrips as prey

Parameters determining the functional and numerical response of Orius insidiosus to the density of second instar larvae of the western flower thrips (WFT), Frankliniella occidentalis, were estimated with the aim to be used in models calculating adequate release ratios of predators vs prey. Especially when the prey population must decrease immediately after predator release and the time to suppression must be short, it is of crucial importance to know whether the functional response reaches a plateau or keeps on increasing with prey density within the range of densities that are realistic for the prey. Such plateaus may arise from constraints on the time budget or from constraints on gut-fullness-associated motivation to attack the prey. Estimates of the plateau for O. insidiosus based on prey handling times by far exceed realistic values. In addition, motivation constraints did not apply, because the level of gut fullness above which attack ceases coincides with gut capacity. This implies that the predation rate keeps on increasing linearly with the square root of the density of WFT-larvae and does not reach a plateau within the realistic range of thrips densities (0–20 WFT per cm2). Such plateaus are likely to occur for smaller-sized predators and smaller-sized stages of the same predator and they may also occur when the prey stage offered has better capacities to escape or resist attack. We argue that the presence and level of plateaus in functional responses are of importance for determining initial predator-to-prey ratios.     

Functional responses of Iphiseius degenerans and Neoseiulus teke (Acari: Phytoseiidae) to changes in the density of the cassava green mite, Mononychellus tanajoa (Acari: Tetranychidae)

The functional responses of protonymph and adult female Iphiseius degenerans and Neoseiulus teke to increasing density of three stages of their prey, the cassava green mite (CGM), Mononychellus tanajoa, were studied on excised cassava leaf discs under laboratory conditions. The responses obtained were predominantly sigmoid type III curves with the highest plateau when both stages of I. degenerans and N. teke were preying on CGM eggs. In all cases, the predation rate of the former species exceeded that of the latter. The empirical data were fitted by four different models. From the models, the attack coefficient (a) and handling time (T _h) were estimated. For a given predator stage (protonymph or adult female), the predator's attack coefficient declines and handling time increases as the prey gets larger. For a given prey stage, the predator's attack coefficient increases and handling time decreases as the predator stage becomes larger.     

Sex-specific developmental plasticity of generalist and specialist predatory mites (Acari: Phytoseiidae) in response to food stress

We studied developmental plasticity under food stress in three female-biased size dimorphic predatory mite species, Phytoseiulus persimilis, Neoseiulus californicus, and Amblyseius andersoni. All three species prey on two-spotted spider mites but differ in the degree of adaptation to this prey. Phytoseiulus persimilis is a specialized spider mite predator, N. californicus is a generalist with a preference for spider mites, and A. andersoni is a broad generalist. Immature predators were offered prey patches of varying density and their survival chances, dispersal tendencies, age and size at maturity measured. Amblyseius andersoni dispersed earlier from and had lower survival chances in low density prey patches than N. californicus and P. persimilis. Age at maturity was not affected by prey density in the generalist A. andersoni, whereas both the specialist P. persimilis and the generalist N. californicus accelerated development at low prey densities. Species-specific plasticity in age at maturity reflects opposite survival strategies when confronted with limited prey: to prematurely leave and search for other food (A. andersoni) or to stay and accelerate development (P. persimilis, N. californicus). In all species, size at maturity was more plastic in females than males, indicating that males incur higher fitness costs from deviations from optimal body size.     

Trait-mediated interactions: influence of prey size, density and experience

1. The role of non-consumptive predator effects in structuring ecological communities has become an important area of study for ecologists. Numerous studies have shown that adaptive changes in prey in response to a predator can improve survival in subsequent encounters with that predator. 2. Prey-mediated changes in the shapes of predators' functional response surfaces determine the qualitative predictions of theoretical models. However, few studies have quantified the effects of adaptive prey responses on the shape of predator functional responses. 3. This study explores how prey density, size and previous predator experience interact to change the functional response curves of different-sized predators. 4. We use a response surface design to determine how previous exposure to small or large odonate predators affected the short-term survival of squirrel tree frog (Hyla squirella) tadpoles across a range of sizes and densities (i.e. the shape of odonate functional response curves). 5. Predator-induced tadpoles in a given size class did not differ in shape, although induction changed tadpole behaviour significantly. Induced tadpoles survived better in lethal encounters with either predator than did similar-sized predator-naive tadpoles. 6. Induction by either predator resulted in increased survival with both predators at a given size. However, different mechanisms led to increased survival for induced tadpoles. Attack rate for the small predators, whereas handling time increased for the large predators.