Cannibalism and Intraguild Predation Among Phytoseiid Mites: Are Aggressiveness and Prey Preference Related to Diet Specialization?

We tested whether specialist and generalist phytoseiid mites differ in aggressiveness and prey choice in cannibalism and intraguild predation. Specialists tested were Galendromus occidentalis, Neoseiulus longispinosus, Phytoseiulus persimilis, and P. macropilis; tested were Amblyseius andersoni, Euseius finlandicus, E. hibisci, Kampimodromus aberrans, Neoseiulus barkeri, N. californicus, N. cucumeris, N. fallacis, and Typhlodromus pyri. Aggressiveness of cannibalistic females against larvae was not related to diet specialization except that highly aggressive species were exclusively generalists. Seldom to moderately cannibalizing species occurred in both generalist and specialist phytoseiids. In contrast to aggressiveness in cannibalism, generalists and specialists differed in aggressiveness in intraguild predation. Adult females of specialists were only slightly aggressive against heterospecific larvae, whereas adult females of all generalists except T. pyri were highly aggressive. Adult females of generalists were able to discriminate between con- and heterospecific larvae and preferentially consumed the latter when given a choice. Adult females of specialists except G. occidentalis showed no preference when given a choice between con- and heterospecific larvae. We conclude that aggressiveness in intraguild predation, species recognition and subsequent preferential consumption of heterospecifics when given a choice is common in generalist but not specialist phytoseiids. We discuss the evolutionary pathways that might have led to the difference between specialists and generalists in species discrimination.     

Intra- and interspecific predation on four life stage groups by the adult females of Metaseiulus occidentalis, Typhlodromus pyri, Neoseiulus fallacis and Amblyseius andersoni

Do adult females of oligophagous species such as Neoseiulus fallacis (Garman) and Metaseiulus occidentalis (Nesbitt) show less intra- and interspecific predation on phytoseiids when other foods are scarce than polyphagous species such as Amblyseius andersoni Chant and Typhlodromus pyri Scheuten? We caged single adult females of each species without food with ten of their own eggs or larvae, with ten eggs or larvae of the other species or with ten nymphs or adult females of M. occidentalis (T. pyri for M. occidentalis). We assessed the ambulatory activity, survival time, egg levels and prey loss in each test. Polyphages (in particular T. pyri) lived longer than oligophages (in particular N. fallacis) without food. The small T. pyri detected its own stages and benefited most by feeding on small active stages of other species. Amblyseius andersoni, the largest mite, fed and gained the most of any species when held with nymphs and female adults. Metaseiulus occidentalis fed on eggs of all four species to enhance survival. The large hyperactive N. fallacis gained the least from these behaviours. Each mite seemed uniquely adapted to survive conditions of scarce prey and these behaviours may explain their roles in phytoseiid mite complexes. Overall, oligophagous adult females fed less and gained less by feeding on phytoseiids than did polyphagous adult females.     

Predators induce egg retention in prey

To prevent predation on their eggs, prey often avoid patches occupied by predators. As a result, they need to delay oviposition until they reach predator-free patches. Because many species allocate energy to egg production in a continuous fashion, it is not clear what kind of mechanism prey use to delay oviposition. We used females of the phytoseiid mite Neoseiulus cucumeris to study these mechanisms. Females were placed in patches with pollen, a food source they use for egg production, and they were exposed to another phytoseiid mite, Iphiseius degenerans, which is an intraguild predator of N. cucumeris juveniles. We found that the oviposition of N. cucumeris females on patches with the predator was lower than on patches without the predator. Cues left by the intraguild predator were not sufficient to elicit such behaviour. Females of N. cucumeris reduced oviposition when exposed to the predator by retaining the egg inside their body, resulting in a lower developmental rate once these eggs were laid. Hence, females are capable of retaining eggs, but the development of these eggs continues inside the mother’s body. In this way, females gain some time to search for less risky oviposition sites.     

Risk assessment of non-target effects caused by releasing two exotic phytoseiid mites in Japan: can an indigenous phytoseiid mite become IG prey?

Two exotic phytoseiid mites, Neoseiulus cucumeris and Amblyseius swirskii, are commercially available in Japan for the control of thrips and other pest insects. As part of a risk assessment of the non-target effects of releasing these two species, we investigated intraguild predation (IGP) between these exotic phytoseiid mites and an indigenous phytoseiid mite Gynaeseius liturivorus, which is promising as an indigenous natural enemy for the control of thrips in Japan. To understand IGP relations between the exotic and indigenous phytoseiid mites after use of the exotic mites for biological control, we investigated IGP between them in the absence of their shared prey. When an IG prey was offered to an IG predator, both exotic and indigenous females consumed the IG prey at all immature stages (egg, larva, protonymph, deutonymph), especially at its larval stages. The propensity for IGP in a no-choice test was measured by the survival time of IG prey corrected using the survival time of thrips offered to the IG predator. There was no significant difference in the propensity for IGP between N. cucumeris and G. liturivorus, but the propensity was significantly higher in A. swirskii than G. liturivorus. The propensity for IGP in a choice test was measured by the prey choice of the IG predator when a conspecific and a heterospecific larva were offered simultaneously as IG prey. Both exotic females consumed the heterospecific larva only. The indigenous female preferentially consumed the heterospecific larva when the heterospecific larva was N. cucumeris, but consumed the conspecific larva when the heterospecific larva was A. swirskii. We concluded that further investigation would be necessary for the exotic mites’ risk assessment, since the propensity for IGP of the two exotic females was similar to or higher than that of the indigenous female in both the no-choice and choice tests.     

Cannibalism and interspecific predation in the phytoseiid mites Phytoseiulus persimilis and Neoseiulus californicus: predation rates and effects on reproduction and juvenile development

We examined intra- and interspecific predation of adult females and immature stages of the generalist Neoseiulus californicus and the specialist Phytoseiulus persimilis. Adult females and immatures of both predators exhibited higher predation rates on larvae than on eggs and protonymphs. N. californicus fed more inter- than intraspecifically. Predation on P. persimilis by N. californicus was more severe than vice versa. P. persimilis had higher predation rates on conspecifics than heterospecifics and was more prone to cannibalism than N. californicus. When provided with phytoseiid prey, P. persimilis suffered higher mortality than N. californicus. When held without food, adult females and protonymphs of N. californicus survived longer than the corresponding stages of P. persimilis. N. californicus females were able to sustain oviposition when preying upon P. persimilis, whereas cannibalizing females did not lay eggs. Females of P. persimilis were not able to sustain oviposition, irrespective of con- or heterospecific prey. Immatures of both predators were able to reach adulthood when provided with either con- or heterospecifics. Juvenile development of N. californicus was shorter with heterospecific vs. conspecific larvae; mortality of P. persimilis immatures was less when feeding on conspecific vs. heterospecific larvae. Different behavioral pattern in intra- and interspecific predation are discussed in regard to their feeding types (generalist vs. specialist).     

有益真绥螨与巴氏新小绥螨的集团内捕食和同类相残作用

【目的】有益真绥螨Euseius utilis是北方地区广泛分布的一种多食性植绥螨,而巴氏新小绥螨Neoseiulus barkeri目前在我国广泛应用于农业害虫的生物防治中。本研究旨在对巴氏新小绥螨在本地的应用进行风险评估及为与有益真绥螨的联合释放提供依据。【方法】本研究在室内通过一系列实验,比较了实验室饲养的巴氏新小绥螨和采自内蒙古农业大学校园的有益真绥螨的雌成螨对同种或异种未成熟螨的捕食量、存活时间及产卵量的差异,检测了有益真绥螨与巴氏新小绥螨两种植绥螨相互之间的攻击强度以及种内和种间的相互作用。【结果】两种植绥螨都难以刺吸同种或异种植绥螨的卵,而对同种或异种植绥螨幼螨的捕食量最大,其次是对若螨。在无共同食物的情况下,有益真绥螨雌成螨对同种植绥螨幼螨的攻击性比对异种植绥螨幼螨的攻击性强(BreslowDay检验:χ~2=13.84,df=1,P0.001),且有益真绥螨对同种植绥螨幼螨的捕食量(9.10±1.65头)高于对异种植绥螨幼螨的捕食量(5.31±1.43头)(T检验:t=5.487,P0.001),巴氏新小绥螨对异种植绥螨幼螨的捕食量(7.48±0.75头)高于对同种植绥螨幼螨的捕食量(4.75±0.58头)(T检验:t=9.110,P0.05)。【结论】有益真绥螨更倾向于捕食同种幼螨而发生同类相残;巴氏新小绥螨更倾向于捕食异种幼螨而发生集团内捕食。     

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.     

Intraguild Interactions Between the Predatory Mites Neoseiulus californicus and Phytoseiulus persimilis

Species at the same trophic level may interact through competition for food, but can also interact through intraguild predation. Intraguild predation is widespread at the second and third trophic level and the effects may cascade down to the plant level. The effects of intraguild predation can be modified by antipredator behaviour in the intraguild prey. We studied intraguild predation and antipredator behaviour in two species of predatory mite, Neoseiulus californicus and Phytoseiulus persimilis, which are both used for control of the two-spotted spider mite in greenhouse and outdoor crops. Using a Y-tube olfactometer, we assessed in particular whether each of the two predators avoids odours emanating from prey patches occupied by the heterospecific predator. Furthermore, we measured the occurrence and rate of intraguild predation of different developmental stages of P. persimilis and N. californicus on bean leaves in absence or in presence of the shared prey. Neither of the two predator species avoided prey patches with the heterospecific competitor, both when inexperienced with the other predator and when experienced with prey patches occupied by the heterospecific predator. Intraguild experiments showed that N. californicus is a potential intraguild predator of P. persimilis. However, P. persimilis did not suffer much from intraguild predation as long as the shared prey was present. This is probably because N. californicus prefers to feed on two-spotted spider mites rather than on its intraguild prey.     

Interspecific competition and predation between immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

Interspecific competition and predation in immature Amblyseius fallacis (Garman), Amblyseius andersoni Chant, Typhlodromus occidentalis (Nesbitt) and Typhlodromus pyri Scheuten were examined in small cages at three egg densities (0, 20 and 80) of two-spotted spider mite, Tetranychus urticae Koch, in the laboratory at 25±1°C,80% RH and 16L: 8D photoperiod. For the six possible between-species comparisons, the large polyphagous A. andersoni always outcompeted the other three predator species, which were either smaller and/or less polyphagous; the small oligophagous T. occidentalis was always eliminated by the other three predator species, which were either larger and/or more polyphagous. The small and polyphagous T. pyri tied with the large and oligophagous A. fallacis. The outcome of the interaction was generally similar at the three prey densities except in (1) the A. fallacis-A. andersoni system where the advantage of A. andersoni over A. fallacis was reduced when 20 or 80 eggs per cage were present at the start of the interaction and (2) the A. fallacis-T. occidentalis system where the advantage of A. fallacis over T. occidentalis increased with prey density. This study indicates that predator size, predator degree of polyphagy and prey density can affect the competitiveness of immature phytoseiids.     

Effects of humidity on eggs and immatures of Neoseiulus fallacis,Amblysieus andersoni,Metaseiulus occidentalis and Typhlodromus pyri (Phytoseiidae): implications for biological control on apple,caneberry, strawberry and hop

The lethal humidity (LH₅₀) responses at 20°C of eggs of two strains of Neoseiulus fallacis (Garman) were 71.6 and 69.7%; of three strains of Amblyseius andersoni (Chant) were 62.9, 62.0 and 62.4% and of one strain each of Typhlodromus pyri Scheuten and Metaseiulus occidentalis Nesbitt were 55.0 and 28.4%, respectively. Eggs of three genetically distinct strains of A. andersoni from Oregon, the Netherlands and Italy did not respond differently from one another nor did eggs of freely hybridizing N. fallacis from Michigan and Oregon. Mortality of larvae through development to early protonymphs at 50% RH, 20°C. was 91.9, 82.3, 46.2 and 31.0% for fed mites and 98.1, 83.2, 67.0 and 89.7% for unfed mites of Oregon strains of N. fallacis, A. andersoni, T. pyri and M. occidentalis, repectively. Fed larvae-protonymphs of M. occidentalis and T. pyri were more tolerant of low humidity than fed larvae-protonymphs of N. fallacis and A. andersoni. Mortality was less for fed than unfed larvae-protonymphs of M. occidentalis and T. pyri, but there were no differences for A. andersoni and N. fallacis. Levels of feeding by predator larvae on T. urticae and cannibalism by phytoseiid protonymphs contributed to species differences. Responses to humidity are discussed in relation to geographic and host plant distributions and biological control by single or mixed species populations of phytoseiids.     

Cannibalism and intraguild predation in Typhlodromus exhilaratus and T. phialatus (Acari: Phytoseiidae) under laboratory conditions

Two species of Phytoseiidae are found in the same agroecosystem: Typhlodromus exhilaratus prevails in vine plots, while T. phialatus prevails in uncultivated surrounding areas. The objective of the present paper was to investigate whether the poor settlement of T. phialatus in vine plots can be explained by intraguild predation of these two species and/or cannibalism. Predatory abilities of the females on larvae and protonymphs were studied under laboratory conditions. A first experiment was conducted with only conspecific or heterospecific phytoseiid prey, in a second experiment Tetranychus urticae eggs were added to the phytoseiid prey. Oviposition, prey consumption, and escape rates of females were recorded. Oviposition and intraguild predation rates were higher for T. exhilaratus than for T. phialatus. Typhlodromus exhilaratus consumed fewer conspecifics than heterospecific phytoseiids, and oviposited when feeding on both diets. Typhlodromus phialatus consumed equal amounts of con- and heterospecifics. Although these two generalist predators belong to the type III defined by McMurtry and Croft (Annual Review of Entomology 42:291–321, 1997), our results suggest that they have different predation behaviour. However, because these results were obtained in experiments where no choice was given between the two phytoseiid species, they are difficult to link to previous studies conducted on the intraguild predation of the Phytoseiidae. The greater voracity and prolificacy of T. exhilaratus could partially explain the poor settlement of T. phialatus in vineyards and the predominance of T. exhilaratus. However, a full understanding of this phenomenon will require the study of other factors, such as susceptibility to pesticides and micro-climatic conditions, as well as the ability to cope with different food sources and host plants.     

Intraspecific competition in immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

Intraspecific competition in immature Amblyseius fallacis, Amblyseius andersoni, Typhlodromus occidentalis and Typhlodromus pyri was examined in the laboratory using small cages at five different predator densities (two, four, eight, 16 and 32) in the absence and presence of prey 100 eggs of two-spotted spider mite, Tetranychus urticae (Koch), at 25 ± 1°C, 80% RH and 16L:8D photoperiod. In the absence of spider mite prey, some individuals of immature phytoseiids showed increased development and surival with increasing predator densities up to certain limits, but none survived to the adult stage, except for a single male each of A. andersoni and A. fallacis who completed development by cannibalizing on conspecifics at a density of 32 predators per cage. In the absence of spider mite prey, the mean immature survival time was independent of the initial predator density, but the variance of survival time increased with predator density. In the presence of prey, the proportion of immatures surviving to adulthood generally decreased with initial predator density and dropped sharply to almost none at the predator density of 32 for A. fallacis, eight for A. andersoni, 16 for T. occidentalis and four for T. pyri. The number of prey consumed per predator during the first day generally decreased with predator density in all four species, as prey available per predator decreased and the competition for food increased with predator density. Our data indicate that scramble competition is operating in these four species. Although cannibalism was occasionally observed, especially after the exhaustion of prey and in the generalist predators such as A. andersoni, the immatures of these phytoseiids were less influenced by the interference of conspecifics than by the increasing difficulty of finding food at high predator densities. The implications of this study for understanding phytoseiid population dynamics and their use in biological control are discussed.     

Effect of pollen supplement on intraguild predatory interactions between two omnivores: The importance of spatial dynamics

Arthropods often engage in complex trophic interactions such as intraguild predation (IGP), true omnivory (i.e., feeding on plants and prey), and apparent competition. Theoretical treatments of the effects of such interactions on herbivore populations have been concerned almost entirely with equilibrium conditions. Yet these interactions are common in non-equilibrium settings such as agroecosystems, where they are likely to have a strong influence on pest populations. We therefore tested short-term effects of IGP and food supplementation on interactions between two predators (the phytoseiid mite Neoseiulus cucumeris and the anthocorid bug Orius laevigatus) and their shared prey, Frankliniella occidentalis, on strawberry plants. All three consumers feed on strawberry pollen, both mites and bugs prey on thrips, and the bug also feeds on the mites (IGP). Strong IGP on mites (IG prey) by the bugs (IG predator) was recorded in structurally-simple arenas. In a more complex setting (whole-plants), however, the intensity of IGP differed among plant structures. Likewise, pollen supplementation reduced both IGP and predation on thrips in a structurally simple setting. In the whole-plant experiment, IGP was more intense on pollen-bearing than pollen-free flowers. The study illustrated how spatial dynamics, generated when consumers track food sources differently in the habitat and possibly when herbivorous and IG prey alter their distribution to escape predation, led to site-specific configuration of interacting populations. The intensity of resulting trophic interactions was weakened by food supplementation and by increased complexity of the habitat.     

Intraguild predation among the predatory mites Amblyseius eharai,Amblyseius cucumeris and Amblyseius barkeri

Predator species with the same prey interact not only by competition for food and space but also by intraguild predation (IGP). The impact of IGP on introduced phytoseiid mites and native species in the context of biological control is a matter of considerable debate. Amblyseius eharai is the dominant native citrus species in central China, while Amblyseius cucumeris and Amblyseius barkeri are candidates for importation. All three species can feed on the spider mite Panonychus citri, which is the main pest in citrus. This study investigated, in the laboratory, possible IGP among these species in the absence and presence of P. citri, respectively. IGP in different densities of shared prey and intraguild (IG) prey was also studied. All three species consumed heterospecific larvae and eggs but not adults, and the IGP rate of larvae was significantly higher than that of eggs in the absence of shared prey. Additionally, the IGP rate of each group was reduced dramatically in the presence of both shared and IG prey when compared to the absence of shared prey. This occurs most likely because the three species prefer to feed on their natural prey P. citri, rather than on IG prey. Our results showed that A. eharai seems to be a more voracious IG predator than A. cucumeris. A. eharai was much more prone to IGP than A. barkeri.     

Oviposition Behavior of Interacting Predatory Mites: Response to the Presence of Con- and Heterospecific Eggs

Oviposition behavior may be affected by the presence of potential future competitors, mates, or predators of offspring. We examined patch choice, oviposition site preference and egg production in the predaceous mites Phytoseiulus persimilis and Neoseiulus californicus (Acari: Phytoseiidae) when given a choice between paired spider mite patches with and without conspecific eggs, with and without heterospecific eggs, and with conspecific or heterospecific eggs. Neoseiulus californicus females had no patch preference and distributed their eggs randomly in all choice situations. This was also the case with P. persimilis females given a choice between patches with and without conspecific eggs and between patches with either con- or heterospecific eggs. Phytoseiulus persimilis females confronted with patches with and without heterospecific eggs preferentially stayed and oviposited in the predator free patches. We discuss the oviposition strategies of P. persimilis and N. californicus with respect to food competition, cannibalism and intraguild predation.     

Larval survival and feeding by immature Metaseiulus occidentalis,Neoseiulus fallacis,Amblyseius andersoni and Typhlodromus pyri on life stage groups of Tetranychus urticae Koch and phytoseiid larvae

When 20 newly hatched larvae either of Metaseiulus occidentalis (Nesbitt), Neoseiulus fallacis (Garman), Amblyseius andersoni Chant or Typhlodromus pyri Scheuten were held in arenas without food at 95% RH and 20°C, the percentages of mites surviving to protonymphs were 5.0, 81.3, 86.3, and 83.8%, respectively. Unfed M. occidentalis larvae starved within 2–3 days, while immatures of the other three species lived up to 12–14 days, with some becoming adults by cannibalizing and/or scavenging. Phytosciid larvae given eggs, larvae/protochrysalis/protonymphs (L/P), deutochrysalis/deutonymphs (D) or teleiochrysalis/female adult (T/A) of Tetranychus urticae Koch, fed at different incidences during 6 h tests. Larvae of T. pyri never fed, but almost all larvae of M. occidentalis fed on eggs and L/Ps and 60–70% of M. occidentalis larvae fed on Ds and T/As. N. fallacis and A. andersoni larvae fed at incidences from 20–75% depending on the stage of spider mite given. Larvae fed more commonly on eggs and L/Ps than Ds and T/As for M. occidentalis and N. fallacis but not A. andersoni. Protonymphs and deutonymphs of all four species, readily fed on T/As after 3 h of exposure, but incidences were higher for A. andersoni and T. pyri. Feeding on phytoseiid larvae by protonymphs and deutonymphs also was more common for A. andersoni and T. pyri. Except for M. occidentalis, deutonymphs fed more than protonymphs on phytoseiid larvae. Results are discussed in relation to individual species life histories and the value of these traits in predicting a species role in a biological control system.     

Absorption and cannibalism: do phytoseiids conserve egg resources when prey densities decline rapidly?

The number of eggs oviposited or left in the opisthosomas of dead mites (total eggs) was assessed for Metaseiulus occidentalis (Nesbitt), Neoseiulus fallacis (Garman), Typhlodromus pyri Scheuten or Amblyseius andersoni Chant when each was caged with either (1) no Tetranychus urticae Koch, (2) only odours of T. urticae, (3) ten eggs of M. occidentalis or (4) ten nymphs of M. occidentalis (T. pyri for M. occidentalis). The total eggs for the no prey versus odour tests did not differ within species; the levels were the greatest for N. fallacis > T. pyri > A. andersoni > M. occidentalis. Among treatments, egg means did not differ for M. occidentalis but they did for N. fallacis and T. pyri and similar trends were seen for A. andersoni. Egg means were usually less for mites held with ten predator nymphs than mites held with ten predator eggs or with no prey. Were adult females with nymphs absorbing rather than ovipositing their eggs or dying with them in their opisthosomas? Activity levels (walking) for adult females were no more for mites held with nymphs versus no food. The data indicated that interference by nymphs was not increasing the energy use of females and thus reducing egg levels. However, tests with ten nymphs, one egg and no adult female had egg losses from nymphal predation that could account for fewer eggs in cage tests. Overall, no evidence for absorption was found. If it occurs, it must be among younger eggs or mites exposed to less rapid prey losses than were the mites tested here; in addition, other stimuli may cause absorption. The total eggs in sticky-tape tests were greatest for N. fallacis > M. occidentalis > T. pyri > A. andersoni. Cage versus stick-tape data differed most for M. occidentalis because of cannibalism. All four mites cannibalized eggs but M. occidentalis did most rapidly and extensively. When starved, it laid all of its eggs before the other three species did. Such behaviours may enhance survival of M. occidentalis when prey become scarce.     

Foraging time and spatial patterns of predation in experimental populations

Summary Responses of the predaceous mites Phytoseiulus persimilis, Typhlodromus (=Metaseiulus) occidentalis, and Amblyseius andersoni to spatial variation in egg density of the phytophagous mite, Tetranychus urticae, were studied in the laboratory.The oligophagous predator P. persimilis showed initially a direct density dependent foraging time allocation and variation in foraging time increased with prey density. With changes in prey density due to predation, predator foraging rates (per hour) decreased with time and density dependent foraging gradually became density independence, because P. persimilis continued to respond to initial prey density, instead of the changing prey density and distribution. The consequent spatial pattern of predation by P. persimilis was density independent, although slopes of predation rate-prey density regressions increased with time.Compared with P. persimilis, the narrowly polyphagous predator T. occidentalis responded relatively slowly to the the presence or absence of prey eggs but not to prey density: the mean and variation of foraging time spent in patches with prey did not differ with prey density, but was significantly greater in patches with prey eggs than in patches without eggs. Prey density and distribution changed only slightly due to predation and overall foraging rates remained more or less constant. The consequent spatial pattern of predation by T. occidentalis was inversely density dependent. As with P. persimilis, slopes of predation rate-prey density regressions increased with time (i.e. the inverse density dependence in T. occidentalis became weaker through time).The broadly polyphagous predator A. andersoni showed density independent foraging time allocation with variation independent of prey density. With changes in prey density over time due to prey depletion, overall foraging rates decreased. The consequent spatial pattern of predation by A. andersoni also changed through time; it initially was inversely density dependent, but soon became density independent.Overall, P. persimilis and T. occidentalis spent more time in prey patches than A. andersoni, suggesting that A. andersoni tended to spend more time moving outside patches. The overall predation rates and searching efficiency were higher in P. persimilis than in A. andersoni and T. occidentalis. Predator reproduction was highest in P. persimilis, lower in T. occidentalis and the lowest A. andersoni.The differences in response to prey distribution among the three predaceous species probably reflect the evolution of these species in environments with different patterns of prey distribution. The degree of polyphagy is a major determinant of the aggregative response, but other attributes such as handling time are also important in other aspects of phytoseiid foraging behavior (e.g. searching efficiency or predation rate).     

Walking,feeding and intraspecific interaction of larvae of Metaseiulus occidentalis,Typhlodromus pyri,Neoseiulus fallacis and Amblyseius andersoni held with and without eggs of Tetranychus urticae

Larvae of Metaseiulus occidentalis (Nesbitt), Typhlodromus pyri Scheuten, Neoseiulus fallacis (Garman) and Amblyseius andersoni Chant exhibited different activity levels when held on apple leaf or on tile arenas and given or not given eggs of Tetranychus urticae Koch and water (tiles only). M. occidentalis larvae held without prey exhibited high levels of walking (includes searching) during 24 hours of evaluation, whereas M. occidentalis larvae held with prey fed quickly and then became less active. Fed larvae of M. occidentalis were less active on leaves than tile. Larvae of T. pyri on leaves had a very low frequency of walking, almost never fed and quickly assumed a resting position during development. While much less active than M. occidentalis, fed and unfed T. pyri larvae walked more on tiles than leaves before resting. Larvae of N. fallacis and A. andersoni fed at low rates and were similarly active on tiles and leaves. Free water increased walking by M. occidentalis, A. andersoni and T. pyri on tile but not N. fallacis. M. occidentalis larvae interacted 5–7 times more often than larvae of the other three species. Cannibalism or scavenging was rarely seen and then only for M. occidentalis larvae. Larvae of all four mites walked, fed and interacted much more in the first 12 hours than the second 12 hours of tests, except unfed M. occidentalis. Unfed M. occidentalis larvae did not molt to protonymphs but unfed larvae of the other three species did. Unfed and fed protonymphs of all four species walked more at 4 hours after molting than larvae at 12–24 hours. Unfed and fed protonymphs of T. pyri or A. andersoni had similar walking frequencies, but unfed protonymphs of N. fallacis were more active than fed ones. Trends in larval activities are discussed relative to the life history of each species.     

Invasion success in communities with reciprocal intraguild predation depends on the stage structure of the resident population

The probability of individuals being targeted as prey often decreases as they grow in size. Such size‐dependent predation risk is very common in systems with intraguild predation (IGP), i.e. when predatory species interact through predation and competition. Theory on IGP predicts that community composition depends on productivity. When recently testing this prediction using a terrestrial experimental system consisting of two phytoseiid mite species, Iphiseius degenerans as the IG‐predator and Neoseiulus cucumeris as the IG‐prey, and pollen (Typha latifolia) as the shared resource, we could not find the predicted community shift. Instead, we observed that IG‐prey excluded IG‐predators when the initial IG‐prey/IG‐predator ratio was high, whereas the opposite held when the initial ratio was low, which is also not predicted by theory. We therefore hypothesized that the existence of vulnerable and invulnerable stages in the two populations could be an important driver of the community composition. To test this, we first demonstrate that IG‐prey adults indeed attacked IG‐predator juveniles in the presence of the shared resource. Second, we show that the invasion capacity of IG‐predators at high productivity levels indeed depended on the structure of resident IG‐prey populations. Third, we further confirmed our hypothesis by mimicking successive invasion events of IG‐predators into an established population of IG‐prey at high productivity levels, which consistently failed. Our results show that the interplay between stage structure of populations and reciprocal intraguild predation is decisive at determining the species composition of communities with intraguild predation.