Phytoseiulus persimilis response to herbivore-induced plant volatiles as a function of mite-days

The predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae), uses plant volatiles (i.e., airborne chemicals) triggered by feeding of their herbivorous prey, Tetranychus urticae (Acari: Tetranychidae), to help locate prey patches. The olfactory response of P. persimilis to prey-infested plants varies in direct relation to the population growth pattern of T. urticae on the plant; P. persimilis responds to plants until the spider mite population feeding on a plant collapses, after which infested plants do not attract predators. It has been suggested that this represents an early enemy-free period for T. urticae before the next generation of females is produced. We hypothesize that the mechanism behind the diminished response of predators is due to extensive leaf damage caused by T. urticae feeding, which reduces the production of volatiles irrespective of the collapse of T. urticae population on the plant. To test this hypothesis we investigated how the response of P. persimilis to prey-infested plants is affected by: 1) initial density of T. urticae, 2) duration of infestation, and 3) corresponding leaf damage due to T. urticae feeding. Specifically, we assessed the response of P. persimilis to plants infested with two T. urticae densities (20 or 40 per plant) after 2, 4, 6, 8, 10, 12 or 14 days. We also measured leaf damage on these plants. We found that predator response to T. urticae-infested plants can be quantified as a function of mite-days, which is a cumulative measure of the standing adult female mite population sampled and summed over time. That is, response to volatiles increased with increasing numbers of T. urticae per plant or with the length of time plant was infested by T. urticae, at least as long at the leaves were green. Predatory mites were significantly attracted to plants that were infested for 2 days with only 20 spider mites. This suggests that the enemy-free period might only provide a limited window of opportunity for T. urticae because relatively low numbers of T. urticae per plant can attract predators. Leaf damage also increased as a function of mite-days until the entire leaf was blanched. T. urticae populations decreased at this time, but predator response to volatiles dropped before the entire leaf was blanched and before the T. urticae population decreased. This result supports our hypothesis that predator response to plant volatiles is linked to and limited by the degree of leaf damage, and that the quantitative response to T. urticae populations occurs only within a range when plant quality has not been severely compromised.     

Evaluation of predatory mites and Acramite for control of twospotted spider mites in strawberries in north central Florida

Greenhouse and field experiments were conducted from 2003 to 2005 to determine the effectiveness of two predatory mite species, Phytoseiulus persimilis Athias-Henriot and Neoseiulus californicus (McGregor), and a reduced-risk miticide, Acramite 50 WP (bifenazate), for control of twospotted spider mite, Tetranychus urticae Koch, in strawberries (Fragaria x ananassa Duchesne). In greenhouse tests, three treatments consisting of releases of P. persimilis, N. californicus, and an untreated control were evaluated. Both species of predatory mites significantly reduced twospotted spider mite numbers below those found in the control during the first 3 wk of evaluation. However, during week 4, twospotted spider mite numbers on the plants treated with P. persimilis increased and did not differ significantly from the control. Field studies used releases of P. persimilis and N. californicus, applications of Acramite, and untreated control plots. Both N. californicus and P. persimilis significantly reduced populations of twospotted spider mite below numbers recorded in the control plots. During the 2003-2004 field season P. persimilis took longer than N. californicus to bring the twospotted spider mite population under control (< 10 mites per leaflet). Acramite was effective in reducing twospotted spider mite populations below 10 mites per leaflet during the 2003-2004 field season but not during the 2004-2005 field season, possibly because of a late application. These findings indicate that N. californicus releases and properly timed Acramite applications are promising options for twospotted spider mite control in strawberries for growers in north Florida and other areas of the southeast.     

Population dynamics of interacting predatory mites,Phytoseiulus persimilis and Neoseiulus californicus,held on detached bean leaves

The success of combined release of the predatory mitesPhytoseiulus persimilis and Neoseiulus californicus insuppression of spider mites may be related to the effects of the interactionsbetween the two predators on their population dynamics. We studied populationgrowth and persistence of the specialist P. persimilis andthe generalist N. californicus reared singly versus rearedin combination after simultaneous and successive predator introductions ondetached bean leaf arenas with abundant prey, Tetranychusurticae, and with diminishing prey. When reared singly with abundantprey, either predator population persisted at high densities to the end of theexperiment. In every predator combination system with abundant prey and variousinitial predator:predator ratios N. californicus displacedP. persimilis. When held singly with diminishing prey, thepopulation of P. persimilis grew initially faster than thepopulation of N. californicus but both species reachedsimilar population peaks. Irrespective whether reared singly or in combination,N. californicus persisted three to five times longer afterprey depletion than did P. persimilis. Regarding thecrucial interactions in the predator combination systems, we conclude thatintraguild predation was a stronger force than food competition and finallyresulted in the displacement of P. persimilis. Previousstudies showed that intraguild predation between the specialist P.persimilis and the generalist N. californicusisstrongly asymmetric favoring the generalist. We discuss the implications ofpotential interactions between P. persimilis andN. californicus to biological control of spider mites.     

Predation preferences and discrimination between con- and heterospecific prey by the phytoseiid mites Phytoseiulus persimilis and Neoseiulus californicus

Discrimination between and predation preference for con- or heterospecific larvae was examined for adult females of P. persimilis and N. californicus in plexiglass cages with and without their primary prey T. urticae. Rates of intra- and interspecific predation on larvae were measured for females held on leaves and provided with excess amounts of spider mites. Females of the generalist N. californicus distinguished con- and heterospecific larvae and preferred to prey upon the latter. Females of the specialist P. persimilis appeared to lack discrimination ability and fed equally on con- and heterospecifics. When spider mites and phytoseiids were offered simultaneously, all P. persimilis females chose to first attack T. urticae, whereas N. californicus females attacked both tetranychids and heterospecific phytoseiids. Females of both predators preyed upon phytoseiid larvae when held on leaves with surplus T. urticae: while P. persimilis fed on both con- and heterospecifics, N. californicus attacked larvae of P. persimilis but avoided cannibalizing larvae. The different behaviors of P. persimilis and N. californicus are discussed with regard to different predation types (generalists vs. specialists) and the possible consequences of mixed release for biological control of spider mites in greenhouses.     

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

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

捕食螨在我国农林害螨生物防治中的应用

捕食螨是农林生产中极具价值的生防作用物。文章对捕食螨在我国农林害螨生物防治中的应用作一综述,内容包括:与释放应用相关的研究(引进与风险评估,饲养、贮存与运输,食性与释放,捕食与繁殖,定殖与扩散,外界影响因子)、几种主要捕食螨(智利小植绥螨Phytoseiulus persimilis、西方盲走螨Typhlodromus occidentalis、伪钝绥螨Amblyseius fallacis、胡瓜钝绥螨A.cucumeris)的应用、释放应用前景与展望等,旨在提高我国的捕食螨研究与应用水平。     

The influence of sublethal deposits of agricultural mineral oil on the functional and numerical responses of <Emphasis Type="Italic">Phytoseiulus persimilis</Emphasis> (Acari: Phytoseiidae) to its prey, <Emphasis Type="Italic">Tetranychus urticae</Emphasis> (Acari: Tetranychidae)

Occasional pesticide application in integrated pest management to at least part of a crop requires that any biological control agents must re-invade previously sprayed areas in order that resurgent pests can be constrained. The ability of the phytoseiid predatory mite Phytoseiulus persimilis to feed on adult two-spotted spider mite (TSSM) Tetranychus urticae on excised leaf discs in both control conditions and in a treatment with a sub lethal residue of agricultural mineral oil (AMO) was assessed. The predator exhibited a Type II functional response with the asymptote significantly higher in the AMO conditions due to the fact that the prey grew slower and reached a smaller size in this treatment. In terms of prey volume eaten, the satiation level of the predator was unchanged by the AMO deposits. The numbers of eggs produced by adult P. persimilis females at densities of 4, 8 and 16 TSSM adult females/disc in the control were significantly higher than those in the AMO treatment, but were similar for the higher density levels, 32 and 64 prey per disc. Thus the functional response in terms of volume of prey eaten explained the numerical response in terms of predator eggs produced. The presence of AMO deposits when the prey were at high density had no effect on predator efficiency (volume eaten) but resulted in a lower intake than that in control conditions when there was a greater distance between prey.     

Spatial distribution of predators and prey affect biological control of twospotted spider mites by Phytoseiulus persimilis in greenhouses

We evaluated the effects of predator release pattern and prey distribution on rate of suppression of the twospotted spider mite, Tetranychus urticae Koch (Acari, Tetranychidae) and visual damage to the ornamental plant, Impatiens wallerana Hook.f., in a greenhouse. Sixteen impatiens plants were arranged in a square and infested with the same total number of spider mites distributed either evenly (equal numbers on all plants) or clumped (divided equally among the 4 central plants), simulating a “hot spot.” The predatory mite, Phytoseiulus persimilis Athias-Henriot, was released at a 1:4 predator:prey ratio based on total spider mites in the experimental unit, but the pattern of release was either even or clumped, which simulated broadcast or point-release strategies, respectively. Nine days after predator release, spider mite populations were reduced in all treatments, but only in the clumped pest-clumped predator treatment were spider mites undetectable. Poorest pest suppression occurred in the clumped spider mite-even predator treatment. Eighteen days after predator release, spider mites were eliminated in all treatments, but a reduction in average plant damage occurred only in treatments in which the predator release pattern matched the spider mite distribution (i.e., even-even or clumped-clumped) with the greatest reduction in the even-even treatment. Results suggest that there is an advantage to releasing predators in “hot spots” provided that the recommended predator:prey ratio is maintained within infested patches. If more uniform predator releases are planned, overall predator numbers need to be kept sufficiently high so that the predator:prey ratio of 1:4 shown to prevent damage on impatiens is achieved in higher-density spider mite patches.     

Vertical dispersal of the two-spotted spider mite Tetranychus urticae, and the predatory mite Phytoseiulus persimilis on dwarf hops

1 The pattern of dispersion within plants of the two-spotted spider mite, Tetranychus urticae, and its predator, the phytoseiid Phytoseiulus persimilis, was studied on the dwarf hop variety First Gold from May to September in 1997 and 1998. 2 Spider mite populations developed on the lower leaves initially but, by late July, as the numbers of mites increased, most were found towards the top of plants. From early August, the numbers of spider mites decreased most rapidly on the upper parts of plants. 3 Where P. persimilis was released, the predator maintained the numbers of T. urticae below those found on non-release plots throughout the season. 4 By early August, the predator’s pattern of dispersion was similar to that of the pest. 5 Predators spread to non-release plots by 20 June in 1997 and 24 July in 1998 and eventually became more numerous than on the plots where they had been released.     

Phytoseiid mites from tropical fruit trees in Bahia State,Brazil (Acari,Phytoseiidae)

The cultivation of tropical fruit trees has grown considerably in the state of Bahia, northeastern Brazil. Some of these have been severely attacked by phytophagous mites, which are usually controlled by the use of chemical pesticides. However, there is today a growing interest for the adoption of less aggressive measures of pest control, as for example the use of predatory mites. Most of the plant-inhabiting predatory mites belong to the family Phytoseiidae. The objective of this paper is to report the phytoseiid species found in an intensive survey conducted on cultivated tropical fruit trees in fifteen localities of the southern coast of Bahia. Measurements of relevant morphological characters are provided for each species, to complement the understanding of the morphological variation of these species. Twenty-nine species of sixteen genera were identified. A key was elaborated to assist in the separation of these species. Fifteen species are reported for the first time in the state, raising to sixty-six the number of species of this family now known from Bahia. Seventy-two percent of the species collected belong to Amblyseiinae, followed by Typhlodrominae (21%) and Phytoseiinae (7%). The most diverse genus was Amblyseius. Amblyseiusoperculatus De Leon was the most frequent and abundant species. Studies should be conducted to evaluate the possible role of the most common predators as control agents of the phytophagous mites co-occurring with them.     

Biological control of twospotted spider mite, <Emphasis Type="Italic">Tetranychus urticae</Emphasis>, with predatory mite, <Emphasis Type="Italic">Neoseiulus californicus</Emphasis>, in strawberries

Greenhouse and field experiments were conducted from 2005 to 2007 to determine the effectiveness of different release times with the predatory mite, Neoseiulus californicus (McGregor), for control of the twospotted spider mite (TSSM), Tetranychus urticae Koch, in strawberries (Fragaria x ananassa Duchesne). The effect of N. californicus releases over time and on development of TSSM populations during a growing season were evaluated. Our hypothesis was that repeated applications of N. californicus, which is currently recommended by biological control companies, might be unnecessary to attain season-long control of TSSM. In greenhouse trials, three treatments consisting of releases of N. californicus at five-day intervals: day 0, day 5, and day 10, and an untreated control were evaluated. The treatment releases significantly reduced TSSM below the control within five days of each release. Neoseiulus californicus significantly reduced TSSM in treatments with high densities (leaflets with ≥ 40 TSSM) below that of treatments with lower densities (leaflets with ≤ 10 TSSM) demonstrating that if released at a predator: prey ratio of 1:10, timing of release does not alter the effectiveness of N. californicus in controlling TSSM. However, we found that if the ratio of predator: prey remains adequate, N. californicus is a more efficient predator at high TSSM densities. Field studies included three treatments consisting of releases of N. californicus at one-month intervals. All treatments significantly reduced TSSM compared with the control plots (no releases). Releases applied early in the season sustained TSSM significantly below those in the control plots for the whole season. Our results indicate that one release of N. californicus is able to sustained control of TSSM in strawberry throughout a growing season if released when TSSM populations are low early in the season in the southeastern United States.     

The Effects of Prey Patchiness, Predator Aggregation, and Mutual Interference on the Functional Response of Phytoseiulus persimilis Feeding on Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae)

The spatial distributions of two-spotted spider mites Tetranychus urticae and their natural enemy, the phytoseiid predator Phytoseiulus persimilis, were studied on six full-grown cucumber plants. Both mite species were very patchily distributed and P. persimilis tended to aggregate on leaves with abundant prey. The effects of non-homogenous distributions and degree of spatial overlap between prey and predators on the per capita predation rate were studied by means of a stage-specific predation model that averages the predation rates over all the local populations inhabiting the individual leaves. The empirical predation rates were compared with predictions assuming random predator search and/or an even distribution of prey. The analysis clearly shows that the ability of the predators to search non-randomly increases their predation rate. On the other hand, the prey may gain if it adopts a more even distribution when its density is low and a more patchy distribution when density increases. Mutual interference between searching predators reduces the predation rate, but the effect is negligible. The stage-specific functional response model was compared with two simpler models without explicit stage structure. Both unstructured models yielded predictions that were quite similar to those of the stage-structured model.     

Comparative toxicity of some acaricides to the predatory mite, Phytoseiulus persimilis and the twospotted spider mite, Tetranychus urticae

The relative toxicity of someacaricides to the predatory mite, Phytoseiulus persimilis and the twospottedspider mite, Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae) wasevaluated in laboratory. Five of theacaricides tested, including bifenazate,acequinocyl, chlorfenapyr, flufenoxuron andfenbutatin oxide, were much less toxic to adultfemales and immatures of P. persimilisthan to those of T. urticae, and adultfemale predators treated with these fiveacaricides produced 84±96% as many eggs as didcontrol females. Etoxazole did not seriouslyaffect the survival and reproduction of adultfemale predators but caused high mortalityrates in eggs and larvae of P.persimilis. Milbemectin and fenazaquin werevery toxic to adult females and immatures ofP. persimilis. Adult female predatorssurvived on a diet of spider mites treated withbifenazate, acequinocyl, chlorfenapyr,flufenoxuron and fenbutatin oxide, and theirfecundity, prey consumption and the sex ratioof the progeny were not substantially affected. Based on the results, bifenazate, acequinocyl,chlorfenapyr, flufenoxuron and fenbutatin oxideappeared to be the promising candidates for usein integrated mite management programs whereP. persimilis is the major naturalenemy.     

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.     

Host range of Neozygites floridana isolates (Zygomycetes: Entomophthorales) to spider mites

Neozygites floridana (Weiser & Muma) (Zygomycetes: Entomophthorales) has been reported infecting naturally at least 18 species of tetranychids worldwide. However, the host range of N. floridana is unknown. Epizootics caused by this pathogen to tetranychid populations indicate that N. floridana has the potential to be used as a biological control agent. However, the virulence and specificity of species and strains of Neozygites need to be assessed in the laboratory to reveal its potential as a biological control agent. N. floridana isolates are currently been investigated in Brazil as biological control agents against the tomato red mite, Tetranychus evansi Baker & Pritchard, and the two-spotted spider mite, Tetranychus urticae Koch. The pathogenicity of five strains of N. floridana obtained from T. urticae, T. evansi and T. ludeni Zacher was assessed against populations of Mononychellus tanajoa (Bondar), Schizotetranychus sacharum Flechtmann & Baker, Tetranychus abacae Baker & Pritchard and Tetranychus armipenis Flechtmann & Baker, in addition to the species from which the fungus was obtained. Mummified mites were placed on leaf discs of the host plant of each tetranychid to promote fungal sporulation, and after 24 h the mites were transferred to the leaf discs. Contamination, infection and mummification were evaluated daily for seven days after confinement. Each isolate was pathogenic to three or four out of the six spider mite species tested. However, except for isolate ESALQ1421, all isolates caused higher levels of infection and significant mummification only to the tetranychid species from which they were collected. None of the isolates was pathogenic to S. sacharum and only one isolate infected T. abacae. Alternative hosts may be important for N. floridana survival in tropical regions where resting spores are rarely found.     

The effect of interspecific mating on sex ratios in the twospotted spider mite and the Banks grass mite (Acarina: Tetranychidae)

Mixed populations of the twospotted spider mite (TSM),Tetranychus urticae (Koch), and the Banks grass mite (BGM),Oligonychus pratensis (Banks), occur on corn and sorghum plants in late summer in the Great Plains. Interspecific matings between these arrhenotokous species occur readily in the laboratory but yield no female offspring. The effect of interspecific mating on female: male sex ratios was measured by examining the F₁ progeny of females that mated with both heterospecific and conspecific males in no-choice situations. TSM females that mated first with BGM males and then with TSM males produced a smaller percentage of female offspring than TSM females that mated only with TSM males (43.1±5.8 and 78.9±2.8% females, respectively). Similarly, BGM females mated with heterospecific males and then with conspecific males produced fewer female offspring than females mated only with BGM males (55.7±5.2 and 77.5±2.5%, respectively). Lower female: male sex ratios were produced also by BGM females that mated with TSM males after first mating with conspecifics (62.4±3.4%). In mixed populations containing males of both species, females also produced lower female: male sex ratios, but these ratios were not as low as expected based on mating propensities and progeny sex ratios observed in no-choice tests. These data suggest that interspecific mating may substantially reduce female fitness in both mite species by reducing the output of female offspring, but in mixed populations this effect is mitigated by unidentified behavioral mechanisms.     

Interactions in a tritrophic acarine predator-prey metapopulation system V: Within-plant dynamics of Phytoseiulus persimilis and Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae)

To investigate the relative contributions of bottom-up (plant condition) and top-down (predatory mites) factors on the dynamics of the two-spotted spider mite (Tetranychus urticae), a series of experiments were conducted in which spider mites and predatory mites were released on bean plants. Plants inoculated with 2, 4, 8, 16, and 32 adult female T. urticae were either left untreated or were inoculated with 3 or 5 adult female predators (Phytoseiulus persimilis) one week after the introduction of spider mites. Plant area, densities of T. urticae and P. persimilis, and plant injury were assessed by weekly sampling. Data were analysed by a combination of statistical methods and a tri-trophic mechanistic simulation model partly parameterised from the current experiments and partly from previous data. The results showed a clear effect of predators on the density of spider mites and on the plant injury they cause. Plant injury increased with the initial number of spider mites and decreased with the initial number of predators. Extinction of T. urticae, followed by extinction of P. persimilis, was the most likely outcome for most initial combinations of prey and predators. Eggs constituted a relatively smaller part of the prey population as plant injury increased and of the predator population as prey density decreased. We did not find statistical evidence of P. persimilis having preference for feeding on T. urticae eggs. The simulation model demonstrated that bottom-up and top-down factors interact synergistically to reduce the density of spider mites. This may have important implications for biological control of spider mites by means of predatory mites.     

Overwintering and effects of autumn habitat manipulation and carbofuran on Neoseiulus fallacis and Tetranychus urticae in peppermint

The percentages of Neoseiulus fallacis (Garman) in four subhabitats of peppermint fields in November were 37% on dead leaves, 45% in debris below dead leaves, 3% in hollow stems and 15% on live foliage. In December, the percentages were 36, 39, 7 and 18%, respectively, for these same sites. The values were 40, 37, 18 and 5% and 6, 80, 14 and 0% for January and February, respectively. By April, 95% of all predators were on new foliage feeding on Tetranychus urticae Koch, while the rest were still in debris. Augmenting plots with debris increased overwintering survival of N. fallacis compared to the natural condition while removal of debris decreased it. An autumn carbofuran (Furadan 4F) treatment completely eliminated overwintering N. fallacis, thus leading to outbreaks of T. urticae the following spring. The results of these overwintering studies are discussed in relation to improved biological control and spider mite management.     

Importance of ambient saturation deficits in an epizootic of the fungus Neozygites floridana in cassava green mites (Mononychellus tanajoa)

The mite-pathogenic fungus Neozygites floridana Fisher (Entomophthorales: Neozygitaceae) is considered to have potential for the biological control of the cassava green mite, Mononychellus tanajoa (Bondar). However, its activity is sporadic and laboratory data suggest a strong dependence on night-time saturation deficits for transmission. We report on an epizootic of this fungus in a mite population in northeastern Brazil. During the epizootic, host populations appeared to be limited by a combination of the pathogen and a predatory mite Neoseiulus idaeus (Acari: Phytoseiidae). When temperatures increased, the epizootic finished and the host population began to grow. Abiotic conditions could not explain the variation in host mortality following pickup of infective propagules in this epizootic. However, night-time saturation did help to explain the variation in transmission from infective cadavers to newly killed hosts. This supports laboratory observations that horizontal transmission between hosts is determined mainly by saturation deficits, while the process of infection is little affected by abiotic conditions. A further field observation was the near-absence of resting spores in dead mites (ca. 0.1% of cadavers), suggesting that the pathogen population was unsuccessful in producing inoculum to infect future M. tanajoa populations. The implications are that this pathogen will only be effective as a biological control agent in periods of high relative humidity, and establishment in new areas may be limited by resting spore formation. This revised version was published online in July 2006 with corrections to the Cover Date.