Plant host associations of Penthaleus species and Halotydeus destructor (Acari: Penthaleidae) and implications for integrated pest management

Integrated pest management programs seek to minimise reliance on pesticides and provide effective long-term control of pests. Cultural control strategies, such as crop rotations, trap and border crops, and weed management, require a thorough understanding of pest host associations. This paper examines the effects of different plant hosts on the persistence and reproduction of blue oat mites, Penthaleus spp., and the redlegged earth mite, Halotydeus destructor (Tucker), which are major agricultural pests in southern Australia. Field and shade-house experiments were conducted testing several crop and plant types. All species survived and reproduced from one mite season to the next when confined to pasture. Canola and a common weed, 'bristly ox-tongue', were suitable hosts for H. destructor and Penthaleus falcatus (Qin and Halliday), whereas Penthaleus sp. x and Penthaleus major (Dugés) failed to persist on these plants. A mixture of wheat and oats sustained P. sp. x and H. destructor, but not P. falcatus or P. major. Lentils were generally a poor host plant for all mite species. These findings show that earth mite species differ in their ability to persist on different plant types, highlighting the importance of distinguishing mite species before implementing control strategies. Results are discussed with respect to cultural control options for the management of these winter pests.     

The biology of Penthaleus species in southeastern Australia

Earth mites are major pests of pastures and crops within southern Australia. While the biology and ecology of the redlegged earth mite (Halotydeus destructor) have been studied extensively, the blue oat mite (Penthaleus major) and its recently discovered relative, Penthaleus falcatus, have received little attention. The distribution of H. destructor, P. major and P. falcatus is described in Victoria, Australia, and the inland borders in central New South Wales are re-described. A new undescribed Penthaleus species identified from mallee samples is recognized, and its restricted distribution in Victoria is mapped. Electrophoretic data and the complete absence of males indicate that all Penthaleus species are obligate thelytokous parthenogens. The three Penthaleus species differ in clonal diversity. All species are pests of pastures and/or crops, however the preferred plant hosts of P. major and P. falcatus differ. Tolerance of a commonly used pesticide (omethoate) differs between the species. P. falcatus shows the highest LC₅₀, and may be responsible for pesticide control failures. Rearing methods for P. major and P. falcatus are described. The large differences between these blue oat mite species point to difficulties in interpreting early published data that failed to distinguish them.     

Diapause and implications for control of Penthaleus species and Halotydeus destructor (Acari: Penthaleidae) in southeastern Australia

Blue oat mites, Penthaleus spp., and redlegged earth mites, Halotydeus destructor (Tucker) are major winter pests of a variety of crops and pastures. In southern Australia earth mites exhibit a facultative egg diapause to survive unfavorable summer conditions. The initiation of diapause egg production in earth mites was investigated using field and shade-house experiments. Species differed in their timing of diapause. H. destructor mainly produced diapausing eggs towards the end of the active mite season in spring, although small numbers were also produced in winter. In contrast, Penthaleus major (Dugés) produced diapause eggs almost immediately after emergence in autumn and continued producing these eggs throughout the season. Penthaleus falcatus (Qin and Halliday) also produced diapause eggs in early winter, although the first appearance of these eggs was slightly later in the season than for P. major. The diapause response of an undescribed species was also somewhat later than in P. major and P. falcatus, but earlier than in H. destructor. Electrophoresis of P. major samples indicated that clones of this parthenogenic species may differ in their timing of diapause egg production, providing another potential selective factor contributing to the maintenance of clonal diversity within this group. The results highlight the importance of determining species composition when devising control strategies for earth mite outbreaks.     

Competitive interactions between two pest species of earth mites, Halotydeus destructor and Penthaleus major (Acarina: Penthaleidae)

Competition was investigated between two pest earth mite species, Penthaleus major Dugés and Halotydeus destructor Tucker, which occur sympatrically and are major pests of southern Australian pastures and crops. Three geographically separate pasture sites within Victoria were sampled monthly. Abundance patterns were similar across sites in that there was a marked increase in number of H. destructor in spring. Field competition plots were set up on pasture. In the first season, H. destructor had a negative effect on P. major but not vice versa. During the summer, diapause and ensuing generation, density-dependent (intraspecific) competition was evident in P. major but not in H. destructor. The interspecific asymmetry was switched in the following season, with P. major dominating most plots and having a negative effect on the reproductive output of H. destructor. These findings indicate that suppression or eradication of one species of earth mite from pastures may result in another species increasing in relative abundance.     

Dispersal patterns of pest earth mites (Acari: Penthaleidae) in pastures and crops

Control methods in pest earth mites and other mites often depend on low dispersal rates, yet there are no experimental estimates of these rates. To rectify this, adult movement rates were estimated in the earth mite Halotydeus destructor Tucker and the winter grain mite, Penthaleus major (Duges), using mark-release-recapture techniques. Mean square dispersal distances were used to estimate diffusion coefficients. In pasture, coefficients were in the range 0.3-1.3 for these species. This suggests that 90% of the population moves < 5-11 m in a 10-d period, or 7-16 m within their adult lifetime. Releases of mites in adjacent pea/wheat crops indicated directional movement toward the more favored pea host. However, there was no directional movement when adjacent plots of peas and lupins were compared, even though lupins are poor hosts. These results indicate that broad border sprays or border culturing will be needed to prevent mite movement from adjacent paddocks.     

Differential levels of mite infestation of wheat and barley in Czech grain stores

Abstract  While mites are able to utilize numerous food sources, the suitability of the food strongly influences population growth. The different suitabilities of various stored agricultural products will thus affect the level of infestation. In this study, we compared field mite infestation rates in two stored cereals: wheat and barley. We analyzed mite abundance, frequency and species composition in samples of grain obtained from 79 selected Czech grain stores. Stored barley seemed to be more vulnerable to mite attack than wheat, as we consistently found more infested samples, more species and higher mean and median mite abundance per sample in barley as compared to wheat. The mean mite abundance per sample were 55 and 506 individuals for wheat and barley, respectively. In barley, 10% of samples exceeded allergen risk threshold (i.e., 1 000 individuals per kg of grain). Altogether, 25 species were identified from approximately 35 000 individuals. The most frequently identified species were the same in wheat and barley, that is, Tydeus interruptus Sig Thor, Acarus siro L., Tarsonemus granarius Lindquist, Lepidoglyphus destructor (Schrank) and Tyrophagus putrescentiae (Schrank). Based on principal components analysis, we found a closer association of T. interruptus , T. putrescentiae , L. destructor and Cheyletus eruditus (Schrank) with barley samples, corresponding to the high frequency and abundance values of these mites. The probable reasons for the higher infestation, especially mite abundance in barley, are discussed in relation to the higher proportion of crushed parts, which may release favorable nutrient sources and amplify the abundance values.     

Predictions of summer diapause in the redlegged earth mite, Halotydeus destructor (Acari: Penthaleidae), in Australia

A prediction for the onset of a summer diapause in the eggs of the redlegged earth mite, Halotydeus destructor, was developed for Australia. In this species diapause eggs pass the summer in the cadavers of adult female mites. Adult female mites were collected for several weeks from pastures in spring at 18 sites in south-western Australia and dissected to determine the timing of the production of diapause eggs. Some sites were sampled for several years between 1990 and 1997. A model was developed to predict the time for onset of diapause. The week at which 90% of eggs were in diapause was predicted best by daylength (80.1% of the variability), then by duration of the long-term plant growing season (10.4%, of variability), leaving 9.5% due to other factors. A single chemical spray in spring 2 weeks before the production of 90% diapause eggs resulted in 99% fewer mites present in autumn 7-8 months later at three sites. The timing of the spring spray was the factor leading to successful control. This model was tested at 17 sites across the whole geographical distribution of the redlegged earth mite in Australia between 1998 and 2001. The observed week of 90% diapause was within 1 week of the predicted week on 81% of occasions, and 2 weeks earlier on 15% of occasions. A database was created for the predicted date of onset of 90% diapause for the whole distribution of the redlegged earth mite in southern Australia on a 10 km(2) grid. Australian farmers are using this for timing a spring spray to control mites in the following autumn.     

Invertebrate pests of canola and their management in Australia: a review

Abstract  In Australia, canola is subject to attack by at least 30 species of invertebrate pests, although the composition of this pest complex can vary between regions. Mites (e.g. the redlegged earth mite Halotydeus destructor and the blue oat mites Penthaleus spp.), lucerne flea ( Sminthurus viridis ) and false wireworms (e.g. the grey false wireworm Isopteron punctatissimus and the bronzed field beetle Adelium brevicorne ) are the major pests threatening the seedling establishment, whereas aphids (the cabbage aphid Brevicoryne brassicae , the turnip aphid Lipaphis erysimi and the green peach aphid Myzus persicae ), the native budworm ( Helicoverpa punctigera ), the diamondback moth ( Plutella xylostella ) and the Rutherglen bug ( Nysius vinitor ) can cause irregular and unpredictable damage to the flowering and podding plants. Current tactics of pest management for canola rely largely on the use of synthetic pesticides, but this single-technology approach is likely to incur negative effects on natural enemies and the risk of pest resistance. Thus, the sustainable production of canola requires integrated pest management (IPM) strategies, in which cultural control, crop resistance and biological control are used as important components, with chemical inputs applied only when absolutely needed to restrict pests from reaching economically damaging densities. Such IPM strategies should be built around a fundamental understanding of pest ecology at both regional and local farm levels and the integration of renewable technologies. Therefore, future research efforts need to be focused on the canola-cropping system, with a particular emphasis on the impact of pest species, natural enemies of the pests, varietal resistance to pests and the spatial ecology of pest species.     

Contrasting structures of plant–mite networks compounded by phytophagous and predatory mite species

Differences in the feeding habits between phytophagous and predatory species can determine distinct ecological interactions between mites and their host plants. Herein, plant–mite networks were constructed using available literature on plant-dwelling mites from Brazilian natural vegetation in order to contrast phytophagous and predatory mite networks. The structural patterns of plant–mite networks were described through network specialization (connectance) and modularity. A total of 187 mite species, 65 host plant species and 646 interactions were recorded in 14 plant–mite networks. Phytophagous networks included 96 mite species, 61 host plants and 277 interactions, whereas predatory networks contained 91 mite species, 54 host plants and 369 interactions. No differences in the species richness of mites and host plants were observed between phytophagous and predatory networks. However, plant–mite networks composed of phytophagous mites showed lower connectance and higher modularity when compared to the predatory mite networks. The present results corroborate the hypothesis that trophic networks are more specialized than commensalistic networks, given that the phytophagous species must deal with plant defenses, in contrast to predatory mites which only inhabit and forage for resources on plants.     

Field associations of first generation densities of the pest mites <Emphasis Type="Italic">Halotydeus destructor</Emphasis> and <Emphasis Type="Italic">Penthaleus major</Emphasis> in pasture

Halotydeus destructor and Penthaleus major are species of earth mite commonly found at high densities in agricultural fields in Australia and other parts of the world. These mites pose a risk to a range of winter crops and pastures when seedlings emerge in autumn. In order to predict likely mite pressure, we investigated whether autumn densities in pastures can be determined from agronomic and environmental field variables. For H. destructor, field densities showed little association with a range of vegetation variables but could largely be explained using the variable field type, with high densities present when fields had mixtures of grass, clover and weeds. For P. major, we found a regional effect. In the region where most data were available, P. major field densities were associated with grass abundance, whereas an association with field type was significant but different to that found for H. destructor. For both species, densities were not associated with rainfall, but there was a weak association with soil moisture capacity. We discuss how these results can help in managing these important pest mites.     

Rapid method for the detection of storage mites in cereals: feasibility of an ELISA based approach

This paper describes the development of rapid immunodiagnostic tests for the detection of storage mite infestations in cereals and cereal products. The study's first phase (proof of concept) involved the production of a species-specific enzyme-linked immunoassay (ELISA) for the flour mite, Acarus siro (L.), a major pest of stored commodities. The specificity of this new assay was assessed against key stored product contaminants (13 species of mites of which three were predatory, five species of insects and five species of fungi) in the presence and absence of grain. The assay was species-specific (no cross-reactivity to other storage contaminants) and was unaffected by the presence of cereal antigens in the extract. In the study's second phase, species- and genera-specific ELISAs were developed for a range of key storage mite pests: the cosmopolitan food mite (Lepidoglyphus destructor), the grocers' itch mite (Glycyphagus domesticus), the grainstack mite (Tyrophagus longior), mites of the Tyrophagus and Glycyphagus generas, and all storage mites. All tests were demonstrably specific to target species or genera, with no cross-reactions observed to other storage pest contaminants or cereals. The final, validation phase, involved a comparative assessment of the species-specific A. siro and the genus-specific Tyrophagus ELISAs with the flotation technique using laboratory and field samples. Both ELISAs were quantitative (0-30 mites per 10 g wheat) and produced good comparative data with the flotation technique (A. siro r(2)=0.91, Tyrophagus spp. r(2)=0.99).     

Hirsutella thompsonii and Metarhizium anisopliae as potential microbial control agents of Varroa destructor,a honey bee parasite

The potential of Hirsutella thompsonii Fisher and Metarhizium anisopliae (Metschinkoff) as biological control agents of the parasitic mite, Varroa destructor Anderson and Trueman was evaluated in the laboratory and in observation hives. In the laboratory, time required for 90% cumulative mortality of mites (LT(90)) was 4.16 (3.98-4.42) days for H. thompsonii and 5.85 (5.48-7.43) days for M. anisopliae at 1.1 x 10(3) conidia mm(-2). At a temperature (34+/-1 degrees C) similar to that of the broodnest in a honey bee colony, Apis mellifera L., H. thompsonii [LC(90)=9.90 x 10(1) (5.86-19.35) conidia mm(-2) at Day 7] and M. anisopliae [LC(90)=7.13 x 10(3) (2.80-23.45) conidia mm(-2) at Day 7] both showed significant virulence against V. destructor. The applications of H. thompsonii to observation hives resulted in significant mortality of mites, and reduction of the number of mites per bee 21 and 42 days post-treatments. The treatments did not significantly affect the mite population in sealed brood. However, the fungus must have persisted because infected mites were still observed [82.97+/-(0.6)%] 42 days post-treatment. In addition, the fungus was found to sporulate on the host. A small percentage [2.86+/-(0.2)%] of dead mites found in the control hives also showed fungal infection, suggesting that adult bees drifted between hives and disseminated the fungus. H. thompsonii was harmless to the honey bees at the concentrations applied and did not have any deleterious effects on the fecundity of the queens. Microbial control with fungal pathogens provides promising new avenues for control of V. destructor and could be a useful component of an integrated pest management program for the honey bee industry.     

Inferring associations among parasitic gamasid mites from census data

Within a community, the abundance of any given species depends in large part on a network of direct and indirect, positive and negative interactions with other species, including shared enemies. In communities where experimental manipulations are often impossible (e.g., parasite communities), census data can be used to evaluate the strength or frequency of positive and negative associations among species. In ectoparasite communities, competitive associations can arise because of limited space or food, but facilitative associations can also exist if one species suppresses host immune defenses. In addition, positive associations among parasites could arise merely due to shared preferences for the same host, without any interaction going on. We used census data from 28 regional surveys of gamasid mites parasitic on small mammals throughout the Palaearctic, to assess how the abundance of individual mite species is influenced by the abundance and diversity of other mite species on the same host. After controlling for several confounding variables, the abundance of individual mite species was generally positively correlated with the combined abundances of all other mite species in the community. This trend was confirmed by meta-analysis of the results obtained for separate mite species. In contrast, there were generally no consistent relationships between the abundance of individual mite species and either the species richness or taxonomic diversity of the community in which they occur. These patterns were independent of mite feeding mode. Our results indicate either that synergistic facilitative interactions among mites increase the host’s susceptibility to further attacks (e.g., via immunosuppression) and lead to different species all having increased abundance on the same host, or that certain characteristics make some host species preferred habitats for many parasite species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Induced responses in clover to an herbaceous mite

Halotydeus destructor feeding on subterranean clover cotyledons can cause severe damage. The mites live on the soil surface and move up onto plants to feed. Foraging behaviour consists of palpating, probing, and feeding with frequent transitions between them. Sustained feeding is made up of a series of short (1-2 min) feeds separated by periods of palpating. The mites tend to feed in aggregations, and are attracted to cotyledons damaged by other mites feeding or by mechanical damage. Mites can distinguish between resistant and susceptible cotyledons within 30 min and resistance is antixenotic due to deterrence. Study of the mechanisms shows this to be induced plant resistance. Several green leaf volatiles are involved in the plant/mite interaction. After feeding commences, 2-E-hexenal is released that at low concentrations is attractive to mites, perhaps causing the feeding aggregations. The wound-induced C(8) compound, 1-octen-3-one, plays a significant role in the deterrence of cotyledons of resistant subterranean clover varieties to H. destructor. Damaged cotyledons of resistant varieties produce more 1-octen-3-one that those of susceptible varieties. Screening for resistance has identified varieties from Italy showing resistance. H. destructor does not occur in Europe. Production of damage-induced volatiles by the resistant plants may have resulted from invasion by herbivores or pathogens, but not from coevolution with these mites. The responses of H. destructor are probably an adaptation to these general plant defensive compounds.     

Host Plant Acceptance by Redlegged Earth Mite, Halotydeus destructor (Tucker) (Acarina: Penthaleidae)

Host plant acceptance behavior of redlegged earth mites (Halotydeus destructor) on cotyledons of subterranean clover (subclover) lines was investigated in the laboratory. Sustained feeding consisted of a series of short bouts of feeding; rather than one long period of feeding. H. destructor preferred to feed at mite-damaged rather than undamaged sites on cotyledons. With time the numbers of mites feeding in aggregations increased, as mites were attracted to damaged patches. Most feeding occurred in aggregations, and mites in such groups benefited by greater weight gain. The sequence of foraging leading to feeding was similar between subclover lines, however, feeding and aggregation activities were markedly reduced on a resistant line. Host plant acceptance occurred during probing but only after some feeding had occurred.     

Spatial associations of insects and mites in stored wheat

The spatial association pattern of insect and mite populations in a steel bin containing stored wheat, Triticum durum Desf., in central Greece, was studied using the Spatial Analysis by Distance IndicEs (SADIE). The monitoring was carried out for 7 mo by using grain trier samples and probe traps. The most abundant insect species were Cryptolestes ferrugineus (Stephens) (Coleoptera: Laemophloeidae) and Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae). For mites, the most abundant species were the phytophagous Lepidoglyphus destructor (Schrank) (Acari: Glycyphagidae) and the predator Blattisocius tarsalis (Berlese) (Mesostigmata: Ascidae). Both for P. interpunctella and C. ferrugineus, trap catches were associated with numbers of individuals in the trier samples, but the overall association index calculated among trap and sample counts was significant only in the 33% of trap-sample pairs of values. Generally, P. interpunctella had the main patch areas in the central part of the bin, with few exceptions, during the entire monitoring period. Similar trends also were noted in the case of C. ferrugineus, which was clearly aggregated in the center of the grain mass. Spatial association maps indicated a stable positive association in the central part of the bin, but in most of the other sampling zones the association was negative. However, distribution of L. destructor, based on trier samples, indicated increased presence in peripheral zones of the grain sampling area. Moreover, B. tarsalis presented the most dispersed distribution among all four species. For each species, the association between two consecutive samplings was significant in the majority of cases, indicating a stable spatial pattern. Finally, B. tarsalis was spatially associated to a higher degree with the insects found rather than with L. desctructor. Moreover, there was no association of insect and mite presence with grain temperature and moisture content. The results of the current study suggest that the coexistence of insects and mites in bulked grain follows a complex pattern, with significant interactions, especially in the case of mite predators, which are spatially associated with insect species.     

Suppressive potential of bean (Phaseolus vulgaris) flour against five species of stored-product mites (Acari: Acarididae)

Previous research has demonstrated that legume proteins have insecticidal activity against stored-product pests, but activity against stored-product mites has not been tested. A study was therefore conducted to explore the potential of bean, Phaseolus vulgaris L., flour as novel botanical acaricide against five species of storage and dust mites: Acarus siro L., Aleuroglyphus ovatus (Troupeau), Caloglyphus redickorzevi (Zachvatkin), Lepidoglyphus destructor (Schrank), and Tyrophagus putrescentiae (Schrank). The effect of wheat, Triticum aestivum L., grain enriched with bean flour to eight concentrations (0, 0.01, 0.1, 0.5, 1, 2.5, 5, and 10%) on population growth initiating from the density of 50 mites per 100 g of wheat was recorded for 21 d under laboratory conditions (grain moisture 14.6% moisture content and 25 degree C in darkness). The enrichment of grain with bean flour suppressed the population growth of all tested species: 0.01% concentration reduced population growth of all tested species to >50% in comparison with the control population. The most sensitive species were A. siro and L. destructor, followed by T. putrescentiae and C. redickorzevi. The least sensitive species was A. ovatus. The terminal (i.e., after 21 d) density of mites positively correlated with bean flour concentration. The suppressive effect of bean flour was not linear but rather asymptotic. The results of this study are discussed in the context of the application of bean flour in integrated control of stored-product mites and the elimination of stored-product mite allergens.     

Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.     

Ecological correlates of body size in gamasid mites parasitic on small mammals: abundance and niche breadth

We studied ecological correlates of body size (abundance and niche breadth) in gamasid mites parasitic on small mammals in 28 regions of the Palearctic. We predicted that smaller species would be characterized by higher abundance than larger species, all else (e.g. host species) being equal. We also predicted that host specificity of mites would decrease (that is, number of host species they use would increase) with an increase in their body size. We focused on mites collected from host bodies that include a) species that feed solely on host’s blood (obligate exclusive haematophages), b) species that feed on both host’s blood and small arthropods (obligate non‐exclusive haematophages), and c) facultative haematophages. We expected that the relationship between body size and abundance and/or host specificity would be more pronounced in obligate exclusively haematophagous mites than for obligate non‐exclusively and facultative haematophagous mites. Across all mite species across regions, mean abundance correlated negatively with body size. The same was true for obligate haematophagous species, but not for facultative haematophages. When mite communities on the same host in a location were considered, the negative body mass–abundance relationship was found in only 3 of 44 communities. Nevertheless, a meta‐analytic (across host species) estimate of the slope of this relationship appeared to be significantly negative. No significant relationship between mite body size and host specificity was found in the analyses across all mite species as well as in obligate exclusive or obligate non‐exclusive haematophages. However, the number of hosts used by facultative haematophagous mites decreased significantly with an increase in their body size. We explain the relationships between morphological (body size) and ecological (abundance and niche breadth) properties of ectoparasites by their interactions with hosts or physical environment.     

Newly isolated bacterial strains belonging to Bacillaceae (Bacillus sp.) and Micrococcaceae accelerate death of the honey bee mite, Varroa destructor (V. jacobsoni), in laboratory assays

Newly isolated bacterial strains belonging to Bacillaceae (Bacillus sp.), Micrococcaceae and three unidentified strains were tested for their pathogenicity against the mite, Varroa destructor. The Bacillus sp. strain and two of the strains belonging to the Micrococcaceae family significantly decreased the time for 50% mortality of the mite population (up to 57%) and hence may be potential control agents. In in vitro bioassay whole cells, extracellular broth and cellular extract of the Bacillus sp. strain effectively killed the mites, suggesting that both endotoxins and exotoxins contributed to the killing.