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.     

Feeding Life Style of Redlegged Earth Mite, Halotydeus Destructor (Acari: Penthaleidae), in pastures and the role of broad-leafed weeds

Redlegged earth mite, Halotydeus destructor, is a major pest of pastures and crops in Australia, and also feeds on lower plants on the soil surface. Feeding behaviour is reviewed, and the role of Arctotheca calendula, capeweed, in determining occurrence and abundance in pastures is investigated. Mites fed more and produced more progeny on Trifolium subterraneum (subclover) than on capeweed in non-choice experiments. In pastures with mixed species three times more mites were feeding on subclover than on capeweed foliage. However, twice as many mites were found on the soil surface under pasture patches consisting mainly of capeweed than of subclover. Patches were selected on the basis of pasture height, irrespective of plant composition. Twice as many mites were found in populations under patches of tall than short pasture. The daytime relative humidity was higher in tall than short patches, and the temperature slightly lower, making tall patches a more favourable niche for these mites. Ninety per cent of H. destructor were on the soil surface, while 10% were feeding on the upper canopy of pasture. The proportion of each population that was feeding was greater on subclover than capeweed, and subclover was a more suitable food. In Australia H. destructor occurs in regions with mixed pasture species, grown in rotation with grain crops. The ability of H. destructor to utilize foliage of a range of plant species of differing suitability for food, while living mostly on the soil surface in niches favourable for survival, has enabled it to become very abundant.     

Antixenotic resistance of subterranean clover cotyledons to redlegged earth mite, Halotydeus destructor

The short-term response of redlegged earth mite, Halotydeus destructor (Tucker) (Acarina: Penthaleidae) to cotyledons of different varieties of subterranean clover (Trifolium subterraneum subsp. subterraneum L.) was assessed by means of paired choice tests, and no-choice tests. H. destructor had lower numbers and fed less on detached cotyledons of resistant than susceptible varieties, yielding a correlation between the numbers of mites and feeding damage to the cotyledons during a three hour test period. For a number of resistant and susceptible varieties, there was a negative correlation between cotyledon deterrence in the three hour choice test and feeding damage to seedling after a two week period. Since the response of the mites to different subclover varieties occurred within three hours, it is concluded that the resistance is based on antixenosis.No evidence was obtained for antifeedant activity in organic solvent extracts from the variety DGI007 (resistant) in comparison with those from the variety Dalkeith (susceptible). Water soluble compounds from DGI007 cotyledons were preferred by mites, in feeding tests in terms of numbers, over those from Dalkeith (susceptible). Squeezed sap from the cotyledons of both varieties showed the same effects on mites as 5% glucose and were more phagostimulatory than water extracts. Mechanically damaged cotyledons of Dalkeith and DGI007 attracted more mites than the undamaged counterparts. The toughness of cotyledons in 17 varieties of T. subterraneum subsp. subterraneum was measured with a manual penetrometer. Results showed a negative correlation between toughness values and mite feeding damage scores (r²=0.752) for all varieties except S3615D (resistant). This implies a likely involvement of epidermal toughness as a contributor in the antixenotic resistance of these varieties. Other mechanisms may be involved in the resistance of S3615D.     

Assays for the effects of volatile compounds from artificially damaged cotyledons of subterranean clover on the redlegged earth mite, Halotydeus destructor

The effects on the redlegged earth mite (Halotydeus destructor) (Acarina: Penthaleidae) of volatile compounds released from artificially damaged cotyledons of subterranean clover (Trifolium subterraneum L.), a host plant, were investigated with different assays. Mites were repelled by the volatile compounds in flask tube and in trimmed cotyledon disc tests. No differences could be shown between different tissue amounts and between resistant and susceptible varieties. When a membrane sachet was used containing either 2-(E)-hexenal, a compound produced by damaged subclover cotyledons, or the total volatile compounds collected from damaged cotyledons of Dalkeith (susceptible) admixed with 1% glucose, mites gathered to low but were deterred by high concentrations of the compounds. Volatile compounds collected from the resistant variety DGI007 were more deterrent than those from the susceptible Dalkeith. Membrane sachets containing 30 p.p.m. of 2-(E)-hexenal and 1% glucose were less preferred than cotyledons of Dalkeith (susceptible) but were not different from DGI007 (resistant). By increasing the glucose concentration to 5%, the membrane sachets with 30 p.p.m. of 2-(E)-hexenal were preferred to cotyledons of either variety. The results indicate that the membrane feeding technique provides a sensitive way of assaying volatile compounds from damaged subclover cotyledons against the mite.     

Avoidance of intraspecific competition via host modification in a grazing, fruit-eating insect

Insects feeding on plants may induce chemical and physical changes in the host plants. Here, we present evidence of host plant modification following an insect attack that may be associated with a reduction in intraspecific competition for food. We demonstrate that feeding by larvae of the cranberry fruitworm, Acrobasis vaccinii, induces a change in fruit colour (from green to red) of cranberry fruits, Vaccinium oxycoccos, that is associated with a significant increase in the concentration of anthocyanin. Host fruit colour affected larval foraging behaviour and food acceptance: significantly more cranberry fruitworm larvae were attracted to, and accepted, green rather than red fruits. Our experiments suggest that fruit reddening also prevents exploitation by conspecific larvae of other green fruits adjacent to the attacked fruit.     

A mycosis of the American lobster, Homarus americanus, caused by Fusarium sp.

Successful inoculations of citrus red mites, Panonychus citri, with the nonoccluded virus were made by feeding healthy mites through membranes on contaminated lemon fruit. Manipulation of the membranes showed that healthy mites pick up the virus from contaminated surface substrates rather than from within plant cells. Electron microscopical studies substantiated that feces of infected mites should be highly infectious since the hindgut contains numerous virus rods.     

Direct and indirect effects of imidacloprid on fecundity and abundance of Eurytetranychus buxi (Acari: Tetranychidae) on boxwoods

Populations of spider mites often reach high levels on urban plants. In many cases, insecticide applications targeting other herbivores trigger outbreaks of spider mites. Recently, elevated populations of spider mites on a diversity of plants in urban landscapes have been associated with applications of imidacloprid, a neonicotinoid insecticide. Imidacloprid has also been linked to increased fecundity in two species of spider mites. In this study, we evaluated the indirect (plant-mediated) and direct impact of imidacloprid on fecundity and longevity of Eurytetranychus buxi Garman (Acari: Tetranychidae), feeding on boxwoods, Buxus sempervirens L. Moreover, we compared the abundance of E. buxi on imidacloprid-treated and untreated boxwoods in the landscape and a greenhouse to determine if changes in the fecundity of mites exposed to imidacloprid were linked to outbreaks of E. buxi. We found that females consuming imidacloprid-treated plants laid more eggs than females feeding on untreated boxwoods, while their longevity remained unchanged. Fecundity was not affected, however, when spider mites were directly sprayed with imidacloprid and consumed foliage of untreated boxwoods. Furthermore, populations of E. buxi were greater on boxwoods treated with imidacloprid in the landscape and greenhouse. On landscape boxwoods, elevated populations of E. buxi persisted into a second year. We also observed general lack of predators of spider mites on treated and untreated boxwoods in the field suggesting that imidacloprid’s eruptive effect on E. buxi stems more from indirect changes in plant quality than from a loss of top-down regulation from E. buxi’s natural enemies.     

Prefeeding and Acceptance Behavior of an Oligophagous Beetle is Dependent on Plant Suitability and Rearing History

Host prefeeding and acceptance behavior of herbivorous insects may strongly differ depending upon the plant species and previous experience. We examined prefeeding behaviors and acceptance of host and non-host plants by the beetle Phaedon cochleariae in populations with different rearing histories using a camera. Suitable host plants evoked short test biting followed by long feeding durations indicating that these first contacts provide sufficient information to stimulate feeding. In contrast, plants of intermediate quality caused prolonged test biting but feeding duration was rather low. When offered unsuitable plants, beetles showed only rare leaf contacts and test biting, demonstrating that test biting behavior is a useful measure of plant suitability. Rearing history had a minor influence on host plant exploitation behavior.     

Humpback whale (Megaptera novaeangliae) and killer whale (Orcinus orca) feeding aggregations for foraging on herring (Clupea harengus) in Northern Norway

Aggregations of predators on food patches have been documented for both terrestrial and marine animals. Here, we documented for the first time, and investigated, non-predatory aggregations occurring between humpback whales (Megaptera novaeangliae) and killer whales (Orcinus orca) while feeding on wintering Norwegian spring spawning herring (Clupea harengus) in Andfjord, northern Norway. Observational data were collected during 109 opportunistic surveys through three seasons 2013–2016. Killer whales were observed feeding on 59 occasions, with one to three humpback whales involved in 47 of these feeding events (79.7%), and there was an increased probability of finding feeding humpback whales when feeding killer whales also were observed. With killer whales identified as the initiating species in 94.4% of the feeding aggregations for which the first species was known, and with humpback whales joining and feeding on the fish ball afterwards, we suggest that humpback whales may benefit more from these aggregations than the opposite.     

Evolutionary adaptation to host plants in a laboratory population of the phytophagous mite Tetranychus urticae Koch

Summary For evolutionary expansion of host range to occur in an herbivore population, genetic variation in ability to survive on and/or accept new hosts must be present. To determine whether a population of the phytophagous mite Tetranychus urticae contained such variation, I established lines from the population on two hosts on which mites initially showed both high juvenile mortality and low acceptance, tomato and broccoli. In less than ten generations, mites from the line kept on each host showed both lower mortality and greater acceptance on it than mites from a control line kept on lima bean, a favorable host for T. urticae. Host acceptance was measured by the proportion of mites attempting to disperse from leaves of the host. The line kept on tomato but not the one kept on broccoli also increased in development rate on its host. These results and those of a similar previous experiment on cucumber indicate that T. urticae populations can adapt to a diversity of initially unfavorable hosts. T. urticae populations therefore should be able to respond to temporal and spatial variation in host availability by adapting to the most abundant hosts.     

Suitability of tomato leaves for larval development of Potato Tuber Moth,Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae)

The potato tuber moth (PTM) is an important pest of the potato plant and its tuber. With the expansion of its geographic range, the PTM could be potential threat to the tomato (Solanum lycopersicum L.), a congeneric species of the potato. To assess that risk, we tested larval feeding and development of PTM on leaves of five cultivated tomato varieties namely Moneymaker, Campari, Ailsa Craig, LA3475, E6203 and one wild tomato variety S. pimpinellifolium. PTM larvae accepted all tested plant leaves and developed into adults. Larval development was fastest on the Ailsa Craig variety. Pupae developed fastest on the Moneymaker variety and slowest on LA3475. Host acceptance and survival were highest on Ailsa Craig and lowest on LA3475. The significantly highest male proportion occurred on LA3475 variety. The study showed that PTM could be a potential threat to tomato cultivation which is rapidly increasing in temperate regions owing to climate change.     

The effect of host plants on Tetranychus evansi, Tetranychus urticae (Acari: Tetranychidae) and on their fungal pathogen Neozygites floridana (Entomophthorales: Neozygitaceae)

In a series of tritrophic-level interaction experiments, the effect of selected host plants of the spider mites, Tetranychus evansi and Tetranychus urticae, on Neozygites floridana was studied by evaluating the attachment of capilliconidia, presence of hyphal bodies in the infected mites, mortality from fungal infection, mummification and sporulation from fungus-killed mite cadavers. Host plants tested for T. evansi were tomato, cherry tomato, eggplant, nightshade, and pepper while host plants tested for T. urticae were strawberry, jack bean, cotton and Gerbera. Oviposition rate of the mites on each plant was determined to infer host plant suitability while host-switching determined antibiosis effect on fungal activity. T. evansi had a high oviposition on eggplant, tomato and nightshade but not on cherry tomato and pepper. T. urticae on jack bean resulted in a higher oviposition than on strawberry, cotton and Gerbera. Attachment of capilliconidia to the T. evansi body, presence of hyphal bodies in infected T. evansi and mortality from fungal infection were significantly higher on pepper, nightshade and tomato. The highest level of T. evansi mummification was observed on tomato. T. evansi cadavers from tomato and eggplant produced more primary conidia than those from cherry tomato, nightshade and pepper. Switching N. floridana infected T. evansi from one of five Solanaceous host plants to tomato had no prominent effect on N. floridana performance. For T. urticae, strawberry and jack bean provided the best N. floridana performance when considering all measured parameters. Strawberry also had the highest primary conidia production. This study shows that performance of N. floridana can vary with host plants and may be an important factor for the development of N. floridana epizootics.     

Cold storage of Halotydeus destructor (Acari: Penthaleidae) for use in experiments

Survival of medium sized nymphal stages of redlegged earth mite Halotydeus destructor (Tucker) (mainly tritonymphs and deutonymphs) stored under low temperature (1.5°C) in sealed plastic boxes remained more than 50% after 12 days of storage, with some mites surviving for up to eight weeks. Adding fresh subclover leaves into the storage box increased the survival rate of mites from 12% to 28%, 19 days after the storage started. Mites stored for two weeks at low temperatures showed feeding activity in a screening experiment similar to mites collected directly from the field. This indicated that cold storage of redlegged earth mite can be used to build up mite numbers for large screening experiments, or to extend the period of availability of mites collected from the field. However, their reproductive ability was greatly reduced after three weeks at low temperature. Thus, care should be taken when using mites for experiments measuring reproduction. The implications of low temperatures for reducing field populations of mites in midwinter are also discussed.     

Artificial feeding of Varroa destructor through a chitosan membrane: a tool for studying the host-microparasite relationship

Rearing pests or parasites of very small size in the absence of their living host is a challenge for behavioural, physiological and pathological studies. For feeding Varroa destructor, an ectoparasitic mite of Apis mellifera, a confinement space with a membrane separating the nutritive solution and the space was designed. The mite measures less than 2 mm and bears a perforating apparatus with a length of 15 μm. The membrane, an essential element of the chamber, has a thickness of 0.1 μm, and is made of chitosan. It closes one face of the individual confinement chamber and allows piercing and the ingestion of the nutritive solution. Factors inducing feeding can be applied on the inner walls or on the membrane. In the particular case of Varroa, the highest percentages of feeding mites are obtained by addition of host haemolymph to the nutritive solution, suggesting the kairomonal role of haemolymph in addition to its nutritional one. The membrane concept can be easily applied to several mites or other micro-pests.     

Host-plant acceptance and performance of Tetranychus urticae (Acari, Tetranychidae)

Abstract: Tetranychus urticae is a serious pest of several crops worldwide. It performs differentially on diverse host‐plant species. Because dispersion is mainly passive, the process of host‐plant selection should be viewed in terms of host‐plant acceptance and not in terms of host finding. The objectives of this study were to investigate: (1) the acceptance of strawberry (S) in comparison with onion (O), leek (L) and parsley (P) by T. urticae; and (2) the suitability of these host plants for the performance of T. urticae, in terms of fecundity and survival. Host‐plant acceptance was measured through females that settled on a test plant or females that left for another host plant. In another experiment, for a duration of 5 days, individual newly mated females were assigned to discs of each host plant and the fecundity, the maximum number of offspring obtained from one female that settled on the disc, the survival and the percentage of females that oviposited at least one egg were analysed. A choice test to measure the preference of females for different host plants was performed. Movements took place towards (S) from the three plants: (O), (L) and (P). Mites significantly moved to other places in the Petri dishes from leek‐origin discs. Dispersal of T. urticae from (S) to other host‐plant discs was similar for all plants. Mean fecundity was higher on (S) than on the other plants. The maximum number of offspring per female was highest on (S), lowest on (L) and (O), and intermediate on (P). Survival of females and percentage of females that laid at least one egg were independent of the host plant. The choice test showed that oviposition was higher on (S) and (P) and very poor on (L) and (O) and that more females settled on (S) and (P). Although mites migrated from parsley to strawberry, the first host plant has a higher acceptance of T. urticae in terms of maximum number of offspring per female than onion and leek, and a lower performance in terms of fecundity than strawberry. Parsley could be a good candidate for an associate plant in a strawberry crop; however, it needs more research under field conditions.     

Influence of sorghum maturity on abundance and damage potential of Banks grass mite,Oligonychus pratensis (Banks)

This research was designed to study the effects of sorghum maturity on abundance and damage potential of Banks grass mite,Oligonychus pratensis (Banks). More mites per plant were recorded on the latest maturing line (M100) and the least on the early maturing lines (60M and CK60). Mite densities on B407 (late maturity) did not differ from any of the lines tested. Mite densities were positively correlated with leaf area. In contrast, mite feeding damage was significantly lower on the late maturing lines, demonstrating that sorghum susceptibility to mite feeding increases as plants reach anthesis and caryopsis.     

Evaluation of Blattisocius dolichus (Acari: Blattisociidae) for biocontrol of root-knot nematode,Meloidogyne incognita (Tylenchida: Heteroderidae)

The root-knot nematode, Meloidogyne incognita Kofoid and White (Tylenchida: Heteroderidae), is one of the most important plant parasitic nematodes attacking many plant roots. In this paper, the predation and biocontrol efficiency of Blattisocius dolichus Ma (Acari: Blattisociidae) on this nematode were studied. The predation rate and the environmental factors affecting predation rate of B. dolichus on second-stage juveniles of M. incognita (Mi-J2) were studied in experimental arenas in plastic dishes. Both female and male mites had greater capability in consuming Mi-J2, and daily predation rate of female and male mites was not less than 35 and 29.6 nematodes within seven days, respectively. Temperature, nematode density and starvation affected B. dolichus predating on Mi-J2, and when offered 300 nematodes per dish, the predation rate of mites starved for 96 h was the highest at 25 °C, with female and male mites consumed 50.5 and 54 nematodes in 24 h, respectively. The biocontrol of B. dolichus against M. incognita was carried out on potted water spinach Ipomoea aquatic in a greenhouse. The water spinach roots were inoculated with 1,000 Mi-J2 ten days after releasing 100, 200, 300, 400 and 500 mites per pot. Compared to the nematodes–alone control, the number of root-knots of all mite treatments and the number of egg masses of the treatments with 300, 400 and 500 mites significantly decreased. Effect of releasing 500 mites on control of the root nematode M. incognita was best among all mite treatments, reduced the root-knots and egg masses 37.1 and 55.1 %, respectively, but no significant difference was observed compared to 400 mites treatment.     

Host Selection by the Herbivorous Mite Polyphagotarsonemus latus (Acari: Tarsonemidae)

This study examined the host-selection ability of the broad mite Polyphagotarsonemus latus (Banks) (Acari: Tarsonemidae). To make long-distance-shifts from one host plant patch to another, broad mites largely depend on phoretic association with whiteflies. However, the host plants of whiteflies and broad mites are not necessarily the same. We determined the host-preference and acceptance of free-moving and phoretic broad mites using two behavioral bioassays. We used a choice test to monitor host selection by free-moving mites. In the case of phoretic mites, we compared their rate of detachment from the phoretic vector Bemisia tabaci placed on leaves taken from various host plants. The suitability of the plant was further determined by monitoring mite’s fecundity and its offspring development. We compared the mites’ responses to young and old cucumber (Cucumis sativus cv. ‘Kfir’) leaves (3rd and 8–9th leaf from the apex, respectively), and two tomato (Solanum lycopersicum cvs. ‘M82’ and ‘Moneymaker’). Free-moving mites of all stages and both sexes preferred young cucumber leaves to old cucumber leaves and preferred young cucumber rather than young tomato leaves, demonstrating for the first time that broad mites are able to choose their host actively. As for phoretic mated females, although eventually most of the mites abandoned the phoretic vector, the rate of detachment from the whitefly vector was host dependent and correlated with the mites’ fitness on the particular host. In general, host preference of phoretic female mites resembled that of the free-moving female. Cues used by mites for host selection remain to be explored.     

Accumulation of photosynthetic pigments in Larix decidua Mill. and Picea abies (L.) Karst. cotyledons treated with 5-aminolevulinic acid under different irradiation

European larch (Larix decidua Mill.) and Norway spruce [Picea abies (L.) Karst.] synthesize chlorophyll (Chl) in darkness. This paper compares Chl accumulation in 14-d-old dark-grown seedlings of L. decidua and P. abies after shortterm (24 h) feeding with 5-aminolevulinic acid (ALA). We used two ALA concentrations (1 and 10 mM) fed to cotyledons of both species in darkness and in continuous light. The dark-grown seedlings of L. decidua accumulated Chl only in trace amounts and the seedlings remained etiolated. In contrast, P. abies seedlings grown in darkness were green and had significantly higher Chl content. After ALA feeding, higher protochlorophyllide (Pchlide) content was observed in L. decidua than in P. abies cotyledons incubated in darkness. Although short-term ALA feeding stimulated the synthesis of Pchlide, Chl content did not change significantly in cotyledons incubated in darkness. The Chl accumulation in cotyledons fed with ALA was similar to the rate of Chl accumulation in the controls. Higher Chl accumulation was reported in control samples after illumination: 86.9% in L. decidua cotyledons and 46.4% in P. abies cotyledons. The Chl content decreased and bleaching occurred in cotyledons incubated with ALA in light due to photooxidation. Analyses of Chlbinding proteins (D1 and LHCIIb) by Western blotting proved differences between Chl biosynthesis in L. decidua and P. abies seedlings in the dark and in the light. No remarkable increase was found in protein accumulation (D1 and LHCIIb) after ALA application. Our results showed interspecific difference in Chl synthesis between two gymnosperms. Shortterm ALA feeding did not stimulate Chl synthesis, thus ALA synthesis was not the rate-limiting step in Chl synthesis in the dark.     

Behavioural response of spider mites (Tetranychus urticae) to induced resistance of cotton plants

ABSTRACT.

• 1 Spider mites (Tetranychus urticae Koch) given a choice between a cotton plant previously damaged by mites and an undamaged control preferentially moved to the control plant.
• 2 This host-choice behaviour was seen in adult female mites but not in immatures.
• 3 Adult females were not found to be significantly more fecund on control plants than on previously damaged plants, but the duration of this choice experiment precluded full assessment of effects on fecundity.
• 4 Mites responded very quickly, choosing previously undamaged plants without contacting or feeding on the test plants. This suggests that an olfactory response to a chemical substance(s) is involved.