Pollen availability for predaceous mites on apple: spatial and temporal heterogeneity

It has been suggested that an abundance of alternate food early in the spring may be critical to the ability of generalist predaceous mites to suppress spider mite pests. One alternate food that is typically very abundant in spring is wind-dispersed pollen. Here we investigate, at several spatial scales, the heterogeneity in the availability of pollen to predaceous mites on apple. We found pollen to be abundant on apple leaves very soon after they opened (>100 grains/cm 2 ), and that the dominant pollen types at this time were wind-dispersed tree pollens (Betulaceae and Pinaceae). We found that most of the spatial variation in pollen abundance occurred at either small spatial scales (within trees) or very large spatial scales (among orchard blocks). Variability among orchards was clearly influenced by the surrounding vegetation, and probably also by the management regime (frequency of mowing). Spatial heterogeneity in pollen availability may affect the build-up of predatory mite populations in the spring, as we found early season abundances of Typhlodromus pyri (Phytoseiidae) and Zetzellia mali (Stigmaeidae) to be better correlated with early season pollen density than with abundance of mite prey (Aculus schlechtendali).     

Quantitative classification of life-style types in predaceous phytoseiid mites

Classification of species into different functional groups based on biological criteria has been a difficult problem in ecology. The difficulty mainly arises because natural classification patterns are not necessarily mutually exclusive. The more group characteristics overlap, the more difficult it is to identify the membership of a species in the overlapping portions of any two groups. In this paper, we present an application of discriminant analysis by creating classification models from life history and morphological data for two specialist and two generalist life-styles type of predaceous phytoseiid mites. Two stages can be distinguished in our method: life-style group membership assignment and trait variable evaluation. We use a Bayesian framework to create a classifier system to locate or assign species within a mixture of trait distributions. The method assumes that a mixture of trait distributions can represent the multiple dimensions of biological data. The mixture is most evident near the boundaries between groups. Because of the complexity of analytical solution, an iterative method is used to estimate the unknown means, variances, and mixing proportion between groups. We also developed a criterion based on information theory to evaluate model performance with different combinations of input variables and different hypotheses. We present a working example of our proposed methods. We apply these methods to the problem of selecting key species for inoculative release and for classical introductions of biological pest control agents.     

Species association among predaceous and phytophagous apple mites (Acari: Eriophyidae, Phytoseiidae, Stigmaeidae, Tetranychidae)

Predator–predator, predator–prey, and prey–prey associations among nine species of mites were studied in a plot of 100 Red Delicious apple (Malus pumila Miller) trees from 1990 to 1997. In 1990, seven-year-old trees were inoculated with Panonychus ulmi (Koch), Tetranychus urticae Koch (Acari: Tetranychidae) or both, and sprayed with azinphosmethyl (alone or plus endosulfan), or nothing. The species Zetzellia mali (Ewing) (Acari: Stigmaeidae), Amblyseius andersoni Chant (Acari: Phytoseiidae), Eotetranychus sp., Bryobia rubrioculus (Scheuten) (Acari: Tetranychidae), and Aculus schlechtendali Nalepa (Acari: Eriophyidae) were already present or immigrated into plots, and Galendromus occidentalis (Nesbitt) and Typhlodromus pyri Scheuten (Acari: Phytoseiidae) were introduced. Yule's V association index was used to measure positive, neutral, or negative interspecific associations for each species pair, because of its robustness with spatially autocorrelated data. We found that pesticide and release treatments did not greatly affect the association results, but there were strong seasonal differences. Predator–predator associations were the strongest and most consistent, showing negative associations in the early and mid seasons, and neutral ones in late season. Negative associations of T. pyri with other predators were the strongest, which is consistent with evidence that this mite can detect other predators on a leaf. Predator–prey seasonal associations were mixed, with some positive and others negative, with most significant associations occurring in the mid season. One prey–prey interaction was positive, again in mid season, most likely because of similar habitat preferences.     

Some predaceous mites [Phytoseiidae] on citrus in the mediterranean region

Seven species of phytoseiid mites were collected from citrus trees in the Mediterranean region during a survey in 1971.Amblyseius stipulatus Athias-Henriot was the most abundant and widespread species in all countries surveyed (Turkey, Greece, Italy, Spain).A. californicus (McGregor) was collected only in Spain;A. potentillae (Garman) andPhytoseiulus persimilis. A.-H. only in Italy.Typhlodromus talbii A.-H. was collected in Greece and Spain,T. athiasae Porath & Swirski in Greece and Turkey, andIphiseius degenerans (Berl.) in Greece and Italy. Living material of all species exceptT. talbii was successfully shipped to California and insectary cultures established. Biologies were studied in the laboratory under 2 food regimes:Tetranychus pacificus McGregor eggs and larvae, and pollen ofMalephora crocea (Jacq.). Pollen was not a satisfactory food forp. persimilis, A. californicus andT. athiasae. Pollen was slightly less satisfactory than tetranychid mites fora. potentillae, but was equally or more favorable than mites forA. stipulatus andI. degenerans. A. stipulatus has become established in southern California.

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Résumé Sept espèces d'acariens de la famille desPhytoseiidae ont été récoltées sur des plants d'agrumes dans la région méditerranéenne au cours d'une étude effectuée en 1971.Amblyseius stipulatus Athias-Henriot est l'espèce la plus abondante et la plus répandue dans tous les pays visités (Turquie, Grèce, Italie, Espagne).A. californicus (McGregor) n'a été trouvé qu'en Espagne.A. potentillae (Garman) etPhytoseiulus persimilis A.-H. n'ont été trouvés qu'en Italie.Typhlodromus talbii. A.-H. a été récolté en Grèce et en Espagne,T. athiasae Porath & Swirski en Grèce et en Turquie, etIphiseius degenerans (Berl.) en Grèce et en Italie. Des spécimens vivants de toutes ces espèces, saufT. talbii, ont été réalisés avec succès en Californie où des élevages en laboratoire ont été réalisés. Ces élevages ont été réalisés avec deux régimes: œufs et larves deTetranychus pacificus, d'une part, pollen deMalephora crocea, d'autre part; ils ont permis de faire les observations biologiques suivantes. Le pollen n'est pas une nourriture satisfaisante pourP. persimilis, A. californicus etT. athiasae. Le pollen convient un peu moins que les tétranyques àA. potentillae; il est aussi favorable ou plus favorable que les tétranyques àA. stipulatus etI. degenerans. A la suite de son introduction dans le sud de la Californie,A. stipulatus semble s'y maintenir.

     

两种捕食螨对苹果园叶螨的控制效果

[目的]以螨治螨是害虫生物防治的一个重要部分,也是果园害虫生态控制的重要措施.通过在苹果园人为释放捕食螨的方法,定量分析加州新小绥螨Neoseiulus californicus(McGregor)和东方钝绥螨Amblyseius orientalis(Ehara)对苹果园2种害螨(苹果全爪螨Panony chusul...     

Rethinking the nature of intraspecific variability and its consequences on species coexistence

Intraspecific variability (IV) has been proposed to explain species coexistence in diverse communities. Assuming, sometimes implicitly, that conspecific individuals can perform differently in the same environment and that IV increases niche overlap, previous studies have found contrasting results regarding the effect of IV on species coexistence. We aim at showing that the large IV observed in data does not mean that conspecific individuals are necessarily different in their response to the environment and that the role of high‐dimensional environmental variation in determining IV has largely remained unexplored in forest plant communities. We first used a simulation experiment where an individual attribute is derived from a high‐dimensional model, representing “perfect knowledge” of individual response to the environment, to illustrate how large observed IV can result from “imperfect knowledge” of the environment. Second, using growth data from clonal Eucalyptus plantations in Brazil, we estimated a major contribution of the environment in determining individual growth. Third, using tree growth data from long‐term tropical forest inventories in French Guiana, Panama and India, we showed that tree growth in tropical forests is structured spatially and that despite a large observed IV at the population level, conspecific individuals perform more similarly locally than compared with heterospecific individuals. As the number of environmental dimensions that are well quantified at fine scale is generally lower than the actual number of dimensions influencing individual attributes, a great part of observed IV might be represented as random variation across individuals when in fact it is environmentally driven. This mis‐representation has important consequences for inference about community dynamics. We emphasize that observed IV does not necessarily impact species coexistence per se but can reveal species response to high‐dimensional environment, which is consistent with niche theory and the observation of the many differences between species in nature.     

Phytophagous and predaceous mites associated with vegetable crops from Riyadh,Saudi Arabia

This study was conducted to investigate phytophagous and predatory mites associated with vegetable plants in Riyadh, Saudi Arabia. Eight phytophagous and 10 predacious mites were collected from 14 species of vegetable crops covering five major production localities. Out of these 18 mite species, 13 species are new to the mite fauna of Saudi Arabia. In addition, the two species, Tenuipalpus punicae and Agistemus exsertus, are reported for the first time on vegetable crops in Saudi Arabia. For each mite species found, notes on host plant association and occurrence period are given. An illustrated key for the identification of the 18 mite species reported in this study is provided and this can be used to improve the IPM programs by applying the local natural predatory mites in controlling mite pests in Saudi Arabia.     

Jerking in predaceous mites (Acari: Phytoseiidae) with emphasis on larvae

Jerking, a behavior involving a pronounced and often repeated lunging of the body, was studied in larvae among seven species of phytoseiid mites, Euseius finlandicus, Galendromus occidentalis, Neoseiulus californicus, Neoseiulus fallacis, Phytoseiulus macropilis, Phytoseiulus persimilis and Typhlodromus pyri, and in the nymphal stages of N. fallacis. This behavior was observed in larvae of six of the seven species and in all active immature stages of N. fallacis. Jerking was usually triggered by direct contact with a con- or heterospecific mite, although jerking occurred occasionally without direct contact in N. fallacis larvae. The larval jerking tendency (the mean fraction of contacts resulting in jerking) of a species was significantly correlated with a tendency to congregate with conspecifics. Congregated larvae often probed their immediate area and each other with the front legs, repeatedly jerking in response to contact with neighboring larvae. Species with larvae having higher jerking tendencies also jerked more repeatedly per jerking event. Among species, there was no consistent relationship between jerking tendency and larval age. Larvae of N. fallacis had a higher tendency to jerk than protonymphs and deutonymphs, and they had both the highest jerking and congregating tendencies among larvae of all species studied. An individual was more likely to jerk when approached by another mite than when it initiated contact, and when observed in N. fallacis larvae that initiated contact, jerking was sometimes aggressive. Jerking N. fallacis larvae experienced fewer and shorter periods of probing by the mouthparts and fewer attacks from cannibalistic adult females than anesthetized larvae that were unable to jerk.     

Washing technique for monitoring mites in apple orchards

A shake-and-wash technique for monitoring the predatory phytoseiid mitesTyphlodromus pyri, Amblyseius finlandicus, and their prey,Panonychus ulmi, Tetranychus urticae andAculus schlechtendali in commercial apple orchards was developed. The removal and recovery of mites from leaves, shoots and spurs is based on agitating the plant material by hand in alcohol and subsequently removing the mites using a separating funnel. The mites are quickly killed and easily washed off the plant material, and are thus well preserved for further study. The technique is more efficient than directly counting the mites on plant material under a dissection microscope and can be easily employed both in the laboratory and in the field.     

Evaluation of astigmatid mites as factitious food for rearing four predaceous phytoseiid mites (Acari: Astigmatina; Phytoseiidae)

Phytoseiids are possibly the most important mites used in biological control and are usually produced using a tritrophic system that, although efficient, is expensive and laborious. Mites of the cohort Astigmatina (Sarcoptiformes) have been used as factitious prey in the mass rearing of phytoseiids and may allow a much simpler production system. This research evaluated the potential of ten Astigmatina species to serve as factitious food sources for Euseius concordis (Chant), Iphiseiodes zuluagai Denmark and Muma, Neoseiulus barkeri Hughes and Neoseiulus californicus McGregor, all phytoseiid species commonly found in different countries. The high fecundity and survival rates obtained suggest that Thyreophagus n. sp. is a suitable prey for rearing N. barkeri and that Austroglycyphagus lukoschusi (Fain) and Blomia tropicalis is suitable for rearing N. californicus. Oviposition by E. concordis was negligible, but survivorship was high on most prey species, suggesting that these species may be useful for maintenance of the predator. I. zuluagai had low fecundity and survival on all the astigmatid species evaluated and none were suitable for its rearing.     

Oviposition and development in predaceous mites fed with artificial and natural diets (Acari: Phytoseiidae)

Oviposition rate and developmental capacity was determined for Amblyseius hibisci (Chant), A. limonicus Garman and McGregor, A. largoensis (Muma), Metaseiulus pomoides Schuster & Pritchard, M. occidentalis (Nesbitt), Typhloseiopsis arboreus (Chant), Typhodromus pyri Scheuten, Phytoseiulus persimilis Athias-Henriot, and Iphiseius degenerans (Berlese) on both artificial and natural diets. Oviposition and development occurred in seven of nine species when fed artificial diet and in all species when fed natural diets (phytophagous mites or pollen). The combined mean oviposition rate (21-d tests) for all species which oviposited on artificial diet was ca. one third the rate achieved by these species when tested on natural diets. Eggs and immature stages derived from oviposition tests with artificial diet showed no abnormalities and developmental times (egg to egg) were similar to those observed on natural diets. Predator cultures maintained on artificial diet showed gradual declines in viability within 2–3 months and only A. hibisci survived longer than 6 months. Adult predators of species which failed to oviposit when fed artificial diet survived longer than adults tested in the absence of food and water. Immature stages of these species failed to develop on artificial diet.

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Ponte et developpement d'acariens predateurs alimentes a partir d'un regime artificiel et d'un regime naturel (Acari: Phytoseiidae)

Résumé Le nombre d'ufs pondus par 7 espèces de Phytoseiidae nourries sur milieu artificiel a varié de 0,1 à 1,2 uf/femelle/jour. Pour l'ensemble des espèces, le nombre moyen d'ufs pondus a été de 0,5 uf/femelle/jour pour une période de ponte moyenne de 18,5 jours.Deux espèces, Phytoseiulus persimilis et Metaseiulus occidentalis nourries sur milieu artificiel, n'ont pas pondu; leurs longévités moyennes ont été respectivement de 11,3 et de 10,3 jours.Avec un régime naturel (acariens phytophages ou pollen), le nombre d'ufs pondus, pour 9 espèces de Phytoseiidae a varié de 0,8 à 3,2 ufs/femelle/jour. Pour l'ensemble des espèces, le nombre moyen d'ufs pondus a été de 1,7 uf/femelle/jour pour une durée moyenne de la période de ponte de 19,0 jours.La viabilité des ufs provenant des expériences de ponte avec régime artificiel était elevée chez toutes les espèces prédatrices et les durées du développement sur milieu artificiel étaient semblables à celles observées avec régime naturel. La viabilité des élevages de prédateurs maintenus sur milieu artificiel a diminué progressivement et la plupart des élevages avaient disparu en moins de 6 mois. Les tentatives d'élevage de P. persimilis et M. occidentalis sur milieu artificiel ont été négatives bien que les prédateurs adultes femelles aient eu une plus grande longévité sur milieu artificiel qu'en absence de nourriture et d'eau.

     

Relatedness among arbuscular mycorrhizal fungi drives plant growth and intraspecific fungal coexistence

Arbuscular mycorrhizal fungi (AMF) form symbioses with most plant species. They are ecologically important determinants of plant growth and diversity. Considerable genetic variation occurs in AMF populations. Thus, plants are exposed to AMF of varying relatedness to each other. Very little is known about either the effects of coexisting AMF on plant growth or which factors influence intraspecific AMF coexistence within roots. No studies have addressed whether the genetics of coexisting AMF, and more specifically their relatedness, influences plant growth and AMF coexistence. Relatedness is expected to influence coexistence between individuals, and it has been suggested that decreasing ability of symbionts to coexist can have negative effects on the growth of the host. We tested the effect of a gradient of AMF genetic relatedness on the growth of two plant species. Increasing relatedness between AMFs lead to markedly greater plant growth (27% biomass increase with closely related compared to distantly related AMF). In one plant species, closely related AMF coexisted in fairly equal proportions but decreasing relatedness lead to a very strong disequilibrium between AMF in roots, indicating much stronger competition. Given the strength of the effects with such a shallow relatedness gradient and the fact that in the field plants are exposed to a steeper gradient, we consider that AMF relatedness can have a strong role in plant growth and the ability of AMF to coexist. We conclude that AMF relatedness is a driver of plant growth and that relatedness is also a strong driver of intraspecific coexistence of these ecologically important symbionts.     

山楂叶螨在苹果不同品种上的生长发育及繁殖

室内试验结果表明 ,山楂叶螨TetrancychusvienensisZacher在苹果不同品种上的生长发育和繁殖有较大的差异。其中在红星上存活率高 ,发育速度快 ,生殖力强 ;而在秦冠上存活率低、发育速度慢、生殖力低。对山楂叶螨种群年结龄构的分析表明 ,卵在种群中所占比例较高 ,而成虫所占比例甚低 ,建议在对其进行防治时选用杀卵力强的药剂     

Comparison of the responses of two predaceous mites,Typhlodromus pyri and Zetzellia mali,to variation in prey density

The ability of a predator to respond to prey density in a patchy habitat has been the focus of much study in biological control systems and elsewhere. Here we look at the response of two species of predatory mite commonly found in commercial apple orchards, Typhlodromus pyri Scheuten and Zetzellia mali Ewing. The two species differ in several characteristics: T. pyri is a more successful biological control agent, is more mobile, has a slightly narrower breadth of diet and prefers the target prey, Panonychus ulmi Koch. We measured resisdence time as a function of prey density, both under field and laboratory conditions. Both predators showed a wide variation in behaviour; however, Z. mali increased residence time in response to the presence of prey, while T. pyri did not show a significant response. Both predators usually left a patch before all prey were consumed. Patterns from the field and laboratory were similar, although residence time was shorter under field conditions.     

An evolutionary explanation of the aggregation model of species coexistence

In ecology, the 'aggregation model of coexistence' provides a powerful concept to explain the unexpectedly high species richness of insects on ephemeral resources like dung pats, fruits, etc. It suggests that females aggregate their eggs across resource patches, which leads to an increased intraspecific competition within occupied patches and a relatively large number of patches that remain unoccupied. This provides competitor-free patches for heterospecifics, facilitating species coexistence. At first glance, deliberately causing competition among the females' own offspring and leaving resources to heterospecific competitors seems altruistic and incompatible with individual fitness maximization, raising the question of how natural selection operates in favour of egg aggregation on ephemeral resource patches. Allee effects that lead to fitness maxima at intermediate egg densities have been suggested, but not yet detected. Using drosophilid flies on decaying fruits as a study system, we demonstrate a hump-shaped relationship between egg density and individual survival probability, with maximum survivorship at intermediate densities. This pattern clearly selects for egg aggregation and resolves the possible conflict between the ecological concept of species coexistence on ephemeral resources and evolutionary theory.     

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

Species coexistence under resource competition with intraspecific and interspecific direct competition in a chemostat

Competition theory has developed separately for direct competition and for exploitative competition. However, the combined effects of the two types of competition on species coexistence remain unclear. To examine how intraspecific and interspecific direct competition contributes to the coexistence of species competing for a single resource, we constructed a chemostat-type resource competition model. With general functions for intraspecific and interspecific direct competition, we derived necessary and sufficient conditions (except for a critical case that rarely occurs in a biological sense) that determine the number of stably coexisting species. From these conditions, we found that the number of coexisting species is determined just by the invasibility of each species into subcommunities with a smaller number of species. In addition, using a combination of rigorous mathematical theory and a simple graphical method, we can demonstrate how the stronger intraspecific direct competition facilitates species invasion, leading to a larger number of coexisting species.     

Species coexistence in resource-limited patterned ecosystems is facilitated by the interplay of spatial self-organisation and intraspecific competition

The exploration of mechanisms that enable species coexistence under competition for a sole limiting resource is widespread across ecology. Two examples of such facilitative processes are intraspecific competition and spatial self-organisation. These processes determine the outcome of competitive dynamics in many resource-limited patterned ecosystems, classical examples of which include dryland vegetation patterns, intertidal mussel beds and subalpine ribbon forests. Previous theoretical investigations have explained coexistence within patterned ecosystems by making strong assumptions on the differences between species (e.g. contrasting dispersal behaviours or different functional responses to resource availability). In this paper, I show that the interplay between the detrimental effects of intraspecific competition and the facilitative nature of self-organisation forms a coexistence mechanism that does not rely on species-specific assumptions and captures coexistence across a wide range of the environmental stress gradient. I use a theoretical model that captures the interactions of two generic consumer species with an explicitly modelled resource to show that coexistence relies on a balance between species' colonisation abilities and their local competitiveness, provided intraspecific competition is sufficiently strong. Crucially, the requirements on species' self-limitation for coexistence to occur differ on opposite ends of the resource input spectrum. For low resource levels, coexistence is facilitated by strong intraspecific dynamics of the species superior in its colonisation abilities, but for larger volumes of resource input, strong intraspecific competition of the locally superior species enables coexistence. Results presented in this paper also highlight the importance of hysteresis in understanding tipping points, in particular extinction events. Finally, the theoretical framework provides insights into spatial species distributions within single patches, supporting verbal hypotheses on coexistence of herbaceous and woody species in dryland vegetation patterns and suggesting potential empirical tests in the context of other patterned ecosystems.     

Patchiness, aggregation, and the coexistence of competitors for ephemeral resources

Abstract. 1. A laboratory experiment was conducted to determine the outcome of competition between two species of carrion-breeding Diptera for food at two levels of resource patchiness.
2. Adults of Fannia howardi Malloch (Muscidae) and Megaselia scalaris Loew (Phoridae) were released into a large population cage and were given access to three 2g sections of pork kidney in one treatment (low patchiness) and to twelve 0.5g sections in the other (high patchiness).
3. Increasing the patchiness of the resource resulted in reduced overlap in the distribution of the two species, and therefore less interspecific competition. However, this did not result in improved survival of the poorer competitor, F.howardi. Both species aggregated their eggs at high patchiness, and therefore suffered from increased levels of intraspecific competition.
4. The experiment shows that increasing patchiness in this way could lead to prolonged coexistence, as has been predicted in other studies.