Nutritional benefits of intraguild predation and cannibalism among generalist and specialist phytoseiid mites

1. Intraguild predation and cannibalism are common among predaceous phytoseiid mites (Acari, Phytoseiidae) but the nutritional benefits gained by these processes are poorly understood. 2. The study reported here addressed the questions of whether cannibalism and intraguild predation provide different nutritional benefits and whether the ability to utilise cannibalism and intraguild predation is linked to the diet specialisation of phytoseiid mites. Specialists tested were Phytoseiulus macropilis, Galendromus occidentalis, and Neoseiulus longispinosus; generalists tested were Amblyseius andersoni, Neoseiulus cucumeris, and Neoseiulus fallacis. 3. All generalists and the specialist P. macropilis were able to complete juvenile development with both con‐ and hetero‐specific prey. Juvenile development of generalists was shorter with heterospecific prey than with conspecific prey, whereas development of the specialist P. macropilis did not differ between prey types. Only a few N. longispinosus and G. occidentalis, both specialists, reached adulthood by cannibalism but none reached adulthood by intraguild predation. 4. All generalists were able to sustain oviposition by intraguild predation. Neoseiulus cucumeris and A. andersoni laid more eggs with heterospecific prey than with conspecific prey, whereas N. fallacis had similar oviposition rates with both prey types. No specialist sustained oviposition by intraguild predation or cannibalism. 5. Overall, generalists gained equal or more nutritional benefits by intraguild predation than by cannibalism and were able to utilise phytoseiid intraguild prey as an alternative food source. Specialists gained equal or more nutritional benefits from cannibalism than from intraguild predation. For specialists, con‐ and hetero‐specific phytoseiids may be considered only a supplemental food.     

Intra- and interspecific predation by adult female Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae) when provisioned with varying densities and ratios of Tetranychus urticae (Acari: Tetranychidae) and phytoseiid larvae

The phytoseiid mites Metaseiulus occidentalis (Nesbitt) and Typhlodromus pyri Schueten are used together and alone as biological control agents against tetranychid pest mites of apple. Their effectiveness as control agents may be impacted by intraguild predation. The effects of prey species and prey density on the rates of inter- and intraspecific predation and oviposition by these two predators were investigated through a series of experiments. Adult female predators were given prey as mixed populations of phytoseiid larvae and larvae of a more preferred species, the spider mite, Tetranychus urticae Koch, at different densities and ratios. Typhlodromus pyri, more of a generalist predator, showed higher rates of predation and cannibalism on phytoseiid immatures at most prey densities and ratios. Manly preference indices indicated that T. pyri switched to feed on phytoseiid larvae at higher prey levels and ratios of T. urticae than M. occidentalis. This greater ability to use phytoseiid larvae as prey may help stabilize T. pyri populations when more preferred prey is unavailable. This may, in part, explain the observed persistence of T. pyri populations when M. occidentalis populations were decreasing in orchard test plots.     

Domatia reduce larval cannibalism in predatory mites

Abstract.  1. Acarodomatia are small structures on the underside of leaves of many plant species, which are mainly inhabited by carnivorous and fungivorous mites.
2. Domatia are thought to protect these mites against adverse environmental conditions and against predation. They are considered as an indirect plant defence; they provide shelter to predators and fungivores and these in turn protect the plants against herbivores and fungi.
3. We studied the possible role of domatia of coffee ( Coffea arabica L.) (Rubiaceae) and sweet pepper ( Capsicum annum L.) (Solanaceae) in reducing cannibalism in the mites inhabiting the domatia. We measured cannibalism of larvae by adults of the predatory mites Iphiseiodes zuluagai Denmark & Muma and Amblyseius herbicolus Chant on coffee leaf discs and of the predatory mite Iphiseius degenerans (Berl.) on sweet pepper leaf. Domatia were closed with glue or left open.
4. Cannibalism in all three species increased when domatia were closed. With I. degenerans , moreover, we found that the previous diet of the cannibal attenuated the effect of domatia on cannibalism.
5. We conclude that domatia can protect young predatory mites against cannibalism by adults and that the diet of cannibals affects the rate of cannibalism.     

非社会性植食昆虫的同种相残行为及其机理

同种相残是指动物杀死并取食同种其他个体的行为,一般被认为是种群密度自我调节的一种方式,常常是影响种群动态和群落结构的重要因素,具有重要的生态学和进化生物学意义。本文从以下几个方面综述了非社会性植食昆虫的同种相残行为及其机理的研究进展：影响同种相残行为发生的因素,同种相残的获利与代价,同种相残与亲子识别及护幼行为的关系,同种相残的进化驱动力等。对植食性昆虫同种相残行为的研究不但具有重要的理论意义,同时,如果我们对同种相残行为发生的生态学、生理学和遗传学机理进行深入研究,或许有助于我们利用相关机理实现植食性昆虫田间种群数量的自我调控,这将对害虫的综合治理产生重要的影响。     

Cannibalism and Interspecific Predation in a Phytoseiid Predator Guild from Cassava Fields in Africa: Evidence from the Laboratory

Interspecific predation and cannibalism are common types of interaction in phytoseiid predator guilds, but the extent and nature of these interactions have not been determined yet in phytoseiid guilds composed of African native and neotropical exotic phytoseiid predators found in cassava habitat in southern Africa. We determined in laboratory experiments the level of cannibalism and interspecific predation among the three phytoseiid mite species Euseius fustis, Iphiseius degenerans, and Typhlodromalus aripo in the absence of food and in the presence of limited or abundant quantities of two food types – Mononychellus tanajoa and maize pollen – commonly found on cassava in Africa. When confined without food, only two T. aripo females laid each two eggs within 5 days, and this species survived longer than I. degenerans and E. fustis. In the presence of con- or hetero-specific larvae or protonymphs, the three species fed more on the former than on the latter, and more on hetero-specifics than on con-specifics. Oviposition rates of the three species did not exceed 0.7 egg/female/day on con- and hetero-specific immatures. Typhlodromalus aripo and E. fustis survived longer on con-specific and hetero-specific larvae and on hetero-specific protonymphs than in the absence of any food, while T. aripo survived longer than the two other species on the same diets. Provision of limited quantity of food decreased interspecific predation rate by I. degenerans and T. aripo, but not by E. fustis, and increased oviposition rate and longevity of all three species. Provision of abundant food, however, eliminated cannibalism by all three species and further reduced interspecific predation rates, but their oviposition and longevity remained relatively unchanged compared with limited food provision. Potential consequences of cannibalism and interspecific predation among phytoseiid mites on cassava for the biological control of M. tanajoa are discussed.     

Phytoseiid mite species composition in Japanese peach orchards estimated using quantitative sequencing

We attempted a population survey of spider mites and phytoseiid mites in Japanese peach orchards with different pesticide practices; however, we had difficulty discriminating phytoseiid mites. To estimate phytoseiid mite species composition, ribosomal gene fragments were amplified from genomic DNA of five phytoseiid mite species using PCR. Cloning and nucleotide sequencing of amplified fragments identified species-specific polymorphic sites. Newly amplified fragments from recombinant plasmids were mixed in various ratios to produce standard DNA template mixtures. After direct sequencing, the signal ratios between two nucleotides at each species-specific polymorphic site were calculated and shown against the corresponding expected ratios. Quadratic regression equations were used to estimate the phytoseiid mite species composition. Results showed that the phytoseiid mite species composition changed during the survey period and varied among study sites.     

Cannibalism in cephalopods

Cannibalism refers to the action of consuming a member of the same species and is common in many taxa. This paper reviews the available literature on cannibalism in cephalopods. All species of the class Cephalopoda are predators and cannibalism is common in most species whose diet has been studied. Cannibalism in cephalopods is density-dependent due to their aggressive predatory and in case of the octopuses territorial nature. It also depends upon local and temporal food availability and of the reproductive season. Cannibalistic behaviour is positively related to the size of both cannibal and victim. It can affect population dynamics of cephalopods in periods of low food availability and/or high population abundance. Cephalopods are generally restricted in their ability to store energy. It is thus assumed that cannibalism is part of a population energy storage strategy enabling cephalopod populations to react to favourable and adverse environmental conditions by increasing and reducing their number. Finally, we propose five orientation points for future research on cannibalism in cephalopods.     

Cannibalizing Harmonia axyridis (Coleoptera: Coccinellidae) larvae use endogenous cues to avoid eating relatives

Cannibalism is widespread among many different organisms, and can have both negative and positive fitness consequences. Avoiding eating relatives can minimize negative fitness consequences of cannibalism. Such avoidance requires kin discrimination, but evidence for this ability among cannibals is limited with little data that address the cues used in such discrimination. We examined whether larvae of the ladybird beetle Harmonia axyridis avoid eating their relatives. We further manipulated environmental factors to create individuals that had similar or dissimilar rearing environments to begin to test for endogenous versus exogenous recognition cues. In our experiments, third-instar larvae were much less likely to cannibalize if they were interacting with a relative. Larvae that did cannibalize kin required more encounters, and significantly delayed cannibalism, compared to larvae cannibalizing unrelated individuals. Acquired cues were less important. Even though the different rearing environment resulted in significantly different phenotypic effects, similarity or dissimilarity of rearing environment had no effect on cannibalism, and there was no interaction between environments and relatedness in cannibalism. We suggest that H. axyridis has a well-developed kin discrimination system, and that kin recognition in this ladybird beetle is based on endogenous rather than exogenous cues. We also argue that these cues reflect either direct or indirect genetic effects on larval phenotypes.     

Immigration of Phytoseiid Mites from Surrounding Uncultivated Areas into a Newly Planted Vineyard

This study reports (1) a faunistic survey of phytoseiid mites observed inside a vine plot and in neighbouring vegetation (other vine plots and uncultivated areas) and (2) dispersal of phytoseiid mites into the plot studied. These data aim to raise some hypotheses concerning natural colonisation of a vineyard by predatory mites. The study was carried out over 3 years (1999, 2000 and 2001) in an experimental plot planted with two cultivars (Grenache and Syrah) and with Sorbus domestica in a framework of agroforestry investigations. Phytoseiid mites were collected in both cultivated and uncultivated areas surrounding the experimental plot, and their dispersal into the plot studied using “aerial” traps. Densities remained quite low compared to previous studies. The main species encountered in the uncultivated areas and in the traps was Typhlodromus phialatus. Despite the low numbers of phytoseiid mites trapped, densities of phytoseiid mites into the vine field increased during 3 years. Typhlodromus phialatus, the species mainly found in the neighbouring uncultivated areas, was rarely found in vineyards. Another morphologically close species was predominant on vines: Typhlodromus exhilaratus. However, Kampimodromus aberrans the main phytoseiid mite species in vineyards of Southern France was not found in the present survey. Hypotheses for this colonisation process are discussed.     

Population survey of phytoseiid mites and spider mites on peach leaves and wild plants in Japanese peach orchard

A population survey of phytoseiid mites and spider mites was conducted on peach leaves and wild plants in Japanese peach orchards having different pesticide practices. The phytoseiid mite species composition on peach leaves and wild plants, as estimated using quantitative sequencing, changed during the survey period. Moreover, it varied among study sites. The phytoseiid mite species compositions were similar between peach leaves and some wild plants, such as Veronica persica, Paederia foetida, Persicaria longiseta, and Oxalis corniculata with larger quantities of phytoseiid mites, especially after mid-summer. A PCR-based method to detect the ribosomal ITS sequences of Tetranychus kanzawai and Panonychus mori from phytoseiid mites was developed. Results showed that Euseius sojaensis (specialized pollen feeder/generalist predator) uses both spider mites as prey in the field.     

Body size,temperature, and seasonal differences in size structure influence the occurrence of cannibalism in larvae of the migratory dragonfly, <Emphasis Type="Italic">Anax junius</Emphasis>

The aim of this study was to test the hypotheses that body size and seasonal differences in temperature and size structure influence cannibalism in larval dragonflies. In the first two experiments, larvae that were either similar or different in size were paired to examine the potential for intra- and intercohort cannibalism. In the third experiment, size structure of an assemblage of larvae and water temperature were manipulated to explore the seasonal dynamics of cannibalism. Cannibalism was common between individuals that differed in body size by one or more instars. Cannibalism also occurred between individuals similar in size but the rate varied across developmental stages. Results suggest that cannibalism may be most common when water temperatures are warm and late-instar larvae are present at high densities. These results highlight the importance of intra- and intercohort cannibalism as factors that can influence the population dynamics of generalist predators.     

Artificial ground shelters for overwintering phytoseiid mites in orchards

Biological control in orchards strongly depends on winter survival of natural enemies, especially in temperate regions. Predacious phytoseiid mites overwinter on trees or on the ground depending on the characteristics of the species. However, the overwintering ecology of phytoseiid mites on the ground is less well known than that of those on trees. We investigated the usefulness of artificial overwintering shelters as a tool for studying the overwintering ecology of phytoseiid mites on the ground. Four kinds of artificial shelter (shading net, felt, cardboard, and urethane foam) were placed on the ground in an apple orchard in Korea. Two dominant phytoseiid species, Neoseiulus makuwa (Ehara) and N. womersleyi (Schicha) (Acari: Phytoseiidae), overwintered in the artificial ground shelters, and numbers were highest in the urethane foam among the four kinds of shelter, and next highest in the shading net. On the other hand, the numbers of phytoseiid mites collected in the ground vegetation plus soil samples under the ground shelters were not significantly different among the five shelter treatments, including the no-shelter control. Our results suggest that artificial overwintering shelters are efficient tools for investigating overwintering ecology of phytoseiid mites on the ground, as well as on trees, in orchards. Furthermore, the artificial shelters would be good sampling units because they are easily formed into identical sizes and can be used almost anywhere in the field with less laborious work. We also discuss some implications about the effects of sheltered structures on the ground on the populations of phytoseiid mites during winter.     

中国植绥螨资源及其生物学研究进展

植绥螨是许多害螨和小型有害昆虫的重要捕食性天敌 ,在农业生产中极具利用价值。我国植绥螨资源相当丰富 ,大量的资源调查工作是在 2 0世纪 70年代末开始的。到目前为止 ,我国已经从不同地区作物上鉴定出植绥螨 2 60余种 ,其中有利用价值的约 2 0种左右。在 2 5～ 3 0℃的适宜温度下 ,植绥螨从卵到羽化为成螨约需 6～ 7d ,而寿命能长达 3 0～ 5 0d。它们对大多数的害螨的功能反应基本成HollingⅡ型。多数成螨为喜湿类型 ,较高的湿度对于孵化和存活尤为重要。在夏季高温和冬季低温来临时进行越夏和越冬 ,以度过不良的环境。植绥螨行两性生殖 ,不交配或交配时间不够 ,不产卵或少产 ,有些种类需要多次交配才能完全产卵。     

Spawning behaviour and acoustic communication in Atlantic cod

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. There are few empirical studies, however, to support this hypothesis. The beaugregory damselfish, Stegastes leucostictus , is a marine teleost that does not actively ventilate its eggs by fanning them. Partial cannibalism is common in this species, but in field studies was found to be unrelated to ration level. Filial cannibalism differed from predation in the pattern of egg eating; filial cannibalism was characterised by a random pattern of egg loss from a clutch rather than an aggregated distribution. Embryos developed quicker and had higher survival rates when they were at low densities and in nest sites where oxygen levels were high, and experimental reduction of oxygen levels increased rates of filial cannibalism. Here I present a hypothesis for filial cannibalism in the beaugregory damselfish; males cannibalise egg clutches in order to reduce clutch density and improve oxygen supply to the remaining embryos. I use a model of filial cannibalism to demonstrate how oxygen mediated cannibalism may be adaptive, and discuss the evolution of filial cannibalism in the beaugregory damselfish and other teleosts.     

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.     

Adaptive filial cannibalism in the beaugregory damselfish

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. There are few empirical studies, however, to support this hypothesis. The beaugregory damselfish, Stegastes leucostictus, is a marine teleost that does not actively ventilate its eggs by fanning them. Partial cannibalism is common in this species, but in field studies was found to be unrelated to ration level. Filial cannibalism differed from predation in the pattern of egg eating; filial cannibalism was characterised by a random pattern of egg loss from a clutch rather than an aggregated distribution. Embryos developed quicker and had higher survival rates when they were at low densities and in nest sites where oxygen levels were high, and experimental reduction of oxygen levels increased rates of filial cannibalism. Here I present a hypothesis for filial cannibalism in the beaugregory damselfish; males cannibalise egg clutches in order to reduce clutch density and improve oxygen supply to the remaining embryos. I use a model of filial cannibalism to demonstrate how oxygen mediated cannibalism may be adaptive, and discuss the evolution of filial cannibalism in the beaugregory damselfish and other teleosts.     

A comparative study of larval cannibalism in three species of ladybird

Abstract. 1. Cannibalism was studied under laboratory conditions in three species of Coccinellidae (Coleoptera): Cycloneda sanguinea , Olla v-nigrum , and Harmonia axyridis . Larval cannibalism varied among species, C. sanguinea  > H. axyridis  ≥  O. v-nigrum .
2. Larvae of all species cannibalised more in response to reduced food availability (14 h starvation daily) than in response to reduced food quality (dry Ephestia eggs).
3. Larvae of H. axyridis cannibalised siblings at lower rates than non-siblings, but larvae of C. sanguinea and O. v-nigrum did not. Rates of cannibalism increased significantly with increasing size disparity among larvae of all three species.
4. Cannibalism in C. sanguinea and H. axyridis , but not in O. v-nigrum , increased with larval density, suggesting that not all attacks on conspecifics are driven by hunger.
5. Costs, not benefits, were the predominant effects of exclusively and partially cannibalistic diets. A diet of conspecific eggs yielded survival equivalent to the Ephestia egg diet for H. axyridis and O. v-nigrum , but developmental time was extended and adults were smaller. Larvae of C. sanguinea had superior survival and faster development on conspecific eggs than on the Ephestia egg diet. Only H. axyridis larvae survived as well on diets comprised exclusively or partially of conspecific larvae as on Ephestia eggs, although they developed more slowly.     

The cannibalistic behaviour of two Gammarus species (Crustacea: Amphipoda)

This study examines the cannibalistic behaviour of the freshwater amphipods Gammarus duebeni celticus Stock & Pinkster, 1970 and G. pulex (L., 1758). In the first experiment, interactions were staged among all combinations of single adult males, single adult females, adults in the precopulatory mate-guarding phase and juveniles. Cannibalism by inter-moult individuals on newly moulted conspecifics occurred in all interaction categories in both species. Gammarus d. celticus , however, were significantly more cannibalistic than G. pulex. Cannibalism between and within sex and size categories (males > females > juveniles) was facilitated by the vulnerability of individuals at moult. Individuals of smaller size categories, however, did not cannibalize newly moulted conspecifics of larger size categories. Males were less cannibalistic on newly moulted females than on newly moulted males and juveniles and, when in the precopulatory condition, appeared to defend females from cannibalistic attacks. In a second experiment, stream conditions were simulated in the laboratory and replicated populations monitored for nine weeks. High levels of cannibalism, and the species and sex differences in cannibalism identified in the first experiment, were confirmed under these heterogeneous conditions. Cannibalism by males on their newly moulted female mating partners, termed 'reversed' sexual cannibalism, was further investigated. When males were deprived of foraging opportunities, cannibalism of precopulatory partners was significantly more frequent. The occurrence of 'reversed' sexual cannibalism is thus interpreted as a conflict between motivation to feed and motivation to mate.     

Ecological and genetic factors influencing evolution of pesticide resistance in tetranychid and phytoseiid mites

Among tetranychid spider mites and their phytoseiid predators, the evolution of pesticide resistance is a common event. In most cases, resistance is based on a single dominant or semidominant gene. However, polygenic, less-stable resistance often develops under laboratory selection. More rapid development of pesticide resistance in spider mites and predatory mites than among other arthropods might partly be due to their arrhenotokous reproduction. For both groups of mites, little study has been done on population genetic factors influencing pesticide resistance. A few studies have focussed on ecological factors. An important ecological factor influencing resistance evolution is the level of immigration of susceptible individuals into treated habitats. Spider mites and predatory mites both tend to reside in treated habitats at high levels and to immigrate at only modest levels from untreated habitats. This favors rapid resistance development. Another factor contributing to rapid resistance evolution in both mite groups is their rapid reproductive rate. A food-limitation factor may limit resistance evolution under field conditions more in predatory mites than spider mites. After treatment by a pesticide, spider mites have an unlimited food source, whereas predatory mites have a decimated food source (their prey), which leads to reduced reproduction, starvation, or migration. Because of the common occurrence of resistance among both mite groups, a strategy of resistance management is often feasible for them. Case histories of IPM where the population dynamics and genetics of pesticide resistance of tetranychid and phytoseiid mites have been considered are discussed. The overall conclusion is that greater understanding of the population genetics and ecology of these species will provide for improved systems of resistance management and IPM.     

Aerial dispersal of phytoseiid mites (Acari: Phytoseiidae): estimating falling speed and dispersal distance of adult females

Aerial dispersal is important to immigration and redistribution of phytoseiid mites that often can provide biological control of spider mite pests. Falling speed of a mite and wind largely determine dispersal distance of such a passively blown organism. A diffusion model of wind-blown phytoseiids could provide insight into their dispersal. To this end, we measured body weights and falling speeds of adult females of 13 phytoseiid and one tetranychid mite species. These data were then incorporated into seed dispersal models (Greene and Johnson, Okubo and Levin) and results were compared to mite dispersal distances in wind tunnel, greenhouse and field. Weights of phytoseiid species ranged from 5.25 to 2l.7 μg; starved mites weighed less than fed mites. Geometric diameters ( d _g ) of idiosomas were correlated to weights. Falling speeds for phytoseiids were 0.39–0.73 m/s, and less than for T. urticae (0.79 m/s) in still air. In some species, active mites had slower falling speeds than inactive (anesthetized) mites indicating that behavior may influence falling. Starved mites had significantly slower falling speeds than fed mites and dispersed farther. Equation-based estimates of falling speed were close to measured ones (2–8% deviation) for some species. There were significant relationships between falling speed and body weight and morphological traits. Greene and Johnson's seed dispersal model provided better fits to dispersal of mites in the wind tunnel, greenhouse and field studies than Okubo and Levin's model. Limits of models in describing mite dispersal distance and applications to IPM are discussed.