Prey-related odor preference of the predatory mites Typhlodromalus manihoti and Typhlodromalus aripo (Acari: Phytoseiidae)

Typhlodromalus manihoti and Typhlodromalus aripo are exotic predators of the cassava green mite Mononychellus tanajoa in Africa. In an earlier paper, we showed that the two predators were attracted to odors from M. tanajoa-infested cassava leaves. In addition to the key prey species, M. tanajoa, two alternative prey mite species, Oligonychus ossypii and Tetranychus urticae also occur in the cassava agroecosystem. Here, we used a Y-tube olfactometer to determine the attraction of the predators to odors from O. gossypii- or T. urticae-infested cassava leaves and their prey-related odor preference. T. aripo but not T. manihoti was slightly attracted to odors from O. gossypii-infested leaves. Both predator species showed a stronger response to odors from cassava leaves infested by M. tanajoa over odors from cassava leaves infested by O. gossypii. Neither predator species was attracted to odors from T. urticae-infested leaves and the predators preferred the odors from M. tanajoa-infested leaves over those from T. urticae-infested leaves. When O. gossypii was present together with M. tanajoa on the same leaves or on different sets of leaves offered together as an odor source the two predators were attracted. In contrast, after mixing non-attractive odors from T. urticae-infested leaves with attractive odors from M. tanajoa-infested leaves, neither T. aripo nor T. manihoti was attracted. Ecological advantages and disadvantages of the predators’ behavior and possible implications for biological control of M. tanajoa are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Patterns, mechanisms and spatial scale of aggregation in generalist and specialist predatory mites (Acari: Phytoseiidae)

Prey species often distribute themselves patchily in their habitats. In response to this spatial variation in prey density, some predator species aggregate in patches of higher prey density. This paper reviews a series of laboratory experiments to demonstrate the patterns of responses by phytoseiid predators (Phytoseiulus persimilis, Typhlodromus occidentalis and Amblyseius andersoni) to spatial variation in the density of their spider mite prey (Tetranychus urticae) and reveal the behavioural mechanisms underlying the observed patterns. In addition, patterns of aggregation were examined at a variety of spatial scales on plants in greenhouses. The patterns, mechanisms and spatial scale of aggregation in three predatory species are discussed in relation to their varying degrees of polyphagy. The results show that a specialist predator species (1) aggregates more strongly than generalist predators, (2) does so not because it finds prey patches of high density more easily but because it remains in these patches longer than generalist predators and (3) tends to aggregate more often at lower levels of spatial scale than generalist predators. It is suggested that these conclusions, based mainly on laboratory studies of a small sample of species, should be tested in the future on a wider selection of specialist and generalist species at different scales in the field. This revised version was published online in November 2006 with corrections to the Cover Date.     

Survival, horizontal distribution, and economics of releasing predatory mites (Acari: Phytoseiidae) using mechanical blowers

The predatory mites Phytoseiulus persimilis and Amblyseius cucumeris are commonly used in commercial greenhouses for augmentative biological control of spider mites and thrips, respectively. One disincentive for growers to use biological control is the cost of manually distributing predators. Therefore, we tested the efficiency of two types of mechanical blowers for dispensing predatory mites. One blower had a metering device to control the flow of material (MBWMD); the other had no metering device (MBWOMD). Predator survival was measured at various distances after dispensing them with blowers. Survival also was compared with a manual-sprinkling method. When blowers were held horizontally, the distribution of predators was uneven for both blowers. For P. persimilis, 77% were found at 1.52 and 2.44 m with the MBWOMD, whereas 75% were located at 2.44, 3.35, and 4.27 m with the MBWMD. For A. cucumeris, 97% were found at 1.52, 2.44, and 3.35 m with the MBWOMD, while 81% landed at 2.44 and 3.35 m with the MBWMD. Percentages of live P. persimilis found using the MBWMD, MBWOMD, and manual-sprinkling were 31 ± 4, 49 ± 3, and 68 ± 8%, respectively. For A. cucumeris, percentages of survival were 61 ± 4, 79 ± 4, and 83 ± 7%, respectively. The MBWOMD dispensed P. persimilis 22 and 44 times faster than the MBWMD and manual-sprinkling, respectively. Mechanical blowers have the advantage of better coverage, substantially less labor, and estimated total costs to growers that are approximately half that of the manual release method.     

Cold-hardiness in the predatory miteAmblyseius potentillae (Acari: Phytoseiidae)

Cold-hardiness was studied in the predatory miteAmblyseius potentillae (Garman) by determining the mortality after exposure to –5°C. Predators reared under diapause-inducing conditions were more cold-resistant than mites that had been kept under long-day conditions. An acclimatization period at 4°C prior to exposure to sub-zero temperatures had a positive effect on the cold-hardiness of the predator, and was remarkable in diapausing mites.Lower temperatures during diapause induction had a positive effect on cold resistance in this predator; mites in which diapause was induced at 15°C were more cold-resistant than mites that had been subjected to short-day conditions at 18 and 20°C.A day/night temperature rhythm did not increase the cold-hardiness of the mite when grown under long-day conditions. Such a rhythm did increase the cold-hardiness of the diapausing predator when given a short acclimatization period, but this effect disappeared after longer acclimatization periods.     

Biosystematics of phytoseiid mites (Acari: Phytoseiidae) associated with cassava

Laboratory crossing experiments indicate that populations of A. limonicus sensu lato collected from cassava (Manihot esculenta Crantz) in Brazil (Cruz das Almas) and Colombia (Palmira and Monteria) are conspecific and different from A. limonicus Garman & McGregor sensu stricto collected in Riverside, California, USA and Jaguariuna, Brazil. A. limonicus s.l. did not cross with A. tenuiscutus, A. rapax or A. limonicus s.s.. A. limonicus s.s. did not cross with A. rapax. Populations of A. aerialis, A. anonymus and A. idaeus from Colombia are conspecific with Brazilian populations of these species.     

Morphological variation and reproductive incompatibility of three coconut-mite-associated populations of predatory mites identified as Neoseiulus paspalivorus (Acari: Phytoseiidae)

Predatory mites identified as Neoseiulus paspalivorus DeLeon (Phytoseiidae) have been considered as agents for classical biological control of the coconut mite, Aceria guerreronis Keifer (Eriophyidae), in Africa and elsewhere. Preliminary identification of geographically distinct populations as belonging to the same species (N. paspalivorus) was based on their morphological similarity. However, laboratory studies recently conducted have shown large differences in feeding behaviors and biological characteristics among individuals collected from three geographic origins: Brazil (South America), Benin and Ghana (West Africa). As morphologically similar specimens do not necessarily belong to the same species, we evaluated under laboratory conditions, reproductive compatibility between the specimens from three geographic locations to ascertain their conspecificity. Morphological measurements were also made to determine whether there is a means of discriminating between them. Inter-population crosses showed complete reproductive isolation between the three geographic populations, but interpopulation discontinuities in morphometric characters were absent. These results indicate that the tested specimens are distinct biological entities despite morphological similarity. Further molecular genetic studies are therefore proposed, including screening for endosymbionts and assessment of genetic differentiation, to determine the cause of reproductive incompatibility and to clarify the taxonomic relationship between those populations.     

Two new predatory mites (Acari: Bdellidae,Phytoseiidae) collected from medicinal plants in West Bengal,India

Two new species of predatory mites, one each of Bdellodes Oudemans (Fam. Bdellidae) and Phytoseius Ribaga (Fam. Phytoseiidae) recorded for the first time from two medicinal plants viz. Ambroma augusta (L.) L.f. (Fam. Sterculiaceae) and Clerodendrum viscosum Vent (Fam. Verbenaceae), respectively, are described in this paper.     

Risk assessment of Cry toxins of Bacillus thuringiensis on the predatory mites Euseius concordis and Neoseiulus californicus (Acari: Phytoseiidae)

Genetically modified plants carrying Cry toxins of Bacillus thuringiensis (Bt) are widely used for pest control. Possible adverse effects as a result of the use of this control technique to non-target organisms is still a concern; however, few studies have addressed the effects of Bt crops on phytoseiid predatory mites. Phytoseiids are important for the natural control of phytophagous mites, but they can also feed on pollen, plant exudates, etc. Thus, phytoseiids may ingest Bt toxins through several pathways. In this paper, we evaluate the direct effect of Bt-toxins by feeding the predators on Bt cell suspensions, on solution of a Bt toxin and the tri-trophic effect by Bt expressed in transgenic plants. We present a method of conducting toxicological tests with Phytoseiidae which can be useful in studies of risk analysis of toxins to be expressed by genetically engineered plants. This method was used to evaluate the potential effect of ingestion of suspensions of Bt (1.25 × 10⁸ spores/ml) and of purified protein Cry1Ia12 (0.006 mg/ml and 0.018 mg/ml) on Euseius concordis, a predatory mite that develops and reproduces best on pollen. The effects of genetically modified Bollgard^® cotton, which carries the Cry1Ac protein, on Neoseiulus californicus, a selective predator that feeds more on spider mites than on pollen or insects, was determined by feeding them with Tetranychus urticae reared in Bollgard^® cotton and on the non-transgenic isoline. When E. concordis was fed with suspension of Bt isolate derived from product Dipel^® PM, no significant effects were detected. Similarly, Cry1Ia12 Bt toxin, at a concentration of 0.006 mg/ml, did not affect E. concordis. At a concentration of 0.018 mg/ml, however, the intake of this protein reduced the reproduction of E. concordis. There were no effects of Bollgard^® cotton on the biological traits and on the predatory capacity of N. californicus. Results indicate that the Cry toxins of B. thuringiensis studied, at the concentrations used in the field or expressed in transgenic plants, should not affect the predatory mites E. concordis and N. californicus.     

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.     

Factors affecting the potential of phytoseiid mites (Acari: Phytoseiidae) as biocontrol agents in North-Italian vineyards

The results of a 5-year study on the relationships between spider mites and their predators in vineyards in Northern Italy are reported. The efficacy of the two predatory mites appeared to be strongly affected by various factors (grape variety, presence of macropredators, climatic condition, interspecific competition, phytoseiid strain). The phytoseiid mitesAmblyseius aberrans (Oud.) andTyphlodromus pyri Scheuten were released at three different density levels in two vineyards (A, B) infested byEotetranychus carpini (Oud.). One strain ofA. aberrans and two strains ofT. pyri were used for the experiments. In vineyard A,Panonychus ulmi (Koch) was recorded in the second and subsequent years of the experiments and became dominant overE. carpini from the third year onwards. The presence of anthocorids in the same vineyard increased the complexity of the system and gave us the possibility of comparing two very different situations. Release of low numbers ofA. aberrans gave a satisfactory control of spider mite populations in both vineyards. These results were even obtained on a variety non preferred by phytoseiids (Merlot) and with the continuous presence of anthocorids (vineyard A). After 5 years,A. aberrans was observed in 53% of the plots in vineyard A and in all plots of the other vineyard (B). Results of experiments in whichT. pyri was released were similar to those obtained in the experiments withA. aberrans, but only in the first year of the study. In vineyard A, theT. pyri populations declined dramatically from the second year onwards; the use of a non-preferred variety (Merlot) and the continuous presence of anthocorids seemed the most important factors causing the decline of predatory mite density. In vineyard B,T. pyri was capable of controlling spider mites even in the second year of the experiments. A very low density of macropredators and a preferred grape variety (Raboso) positively affected control. The density ofT. pyri in vineyard B decreased at the end of the second year because of adverse climatic conditions (high temperatures in combination with a low relative humidity). The decrease ofT. pyri allowed the displacement of this species byA. aberrans in all plots of vineyard B. It was also shown that the twoT. pyri strains differed in their efficacy to control spider mites. The research was partially supported by a grant from Regione Veneto (“Lotta biologica e integrata nel controllo di insetti ed acari dannosi”). The general lines of the research were planned by C. Duso and the most relevant part of the experiments was carried out by the same author. C. Pasqualetto contributed to the experimental work especially during 1989 and 1990.     

Different colonization patterns of phytophagous and predatory mites (Acari: Tetranychidae, Phytoseiidae) on three grape varieties: a case study

In a vineyard having three varieties of grape (Merlot, Trebbiano and Garganega) differently colonized by two phytoseiid species,Typhlodromus pyri Scheuten andAmblyseius andersoni (Chant), the dynamics of mite populations were monitored over 5 years (1989–1993) in order to study their colonization, interspecific competition and the control of spider mites, i.e.Panonychus ulmi (Koch). These aspects were also investigated by releasingT. pyri, A. andersoni andAmblyseius aberrans (Oudemans) on some of the above varieties. In most of the experimental years (1989–1992), selective pesticides were used in order to allow a successful release of phytoseiids, in particularA. aberrans. The use of non-selective insecticides was re-established during 1993 in order to test its effect on the new mite communities originating from 1989 onwards. In the first years of the experiments an apparent relationship between grape variety and phytoseiid species was observed: in the control plots,A. andersoni occurred on Merlot whereT. pyri was rare, while the latter species was largely dominant overA. andersoni on Trebbiano and Garganega.Panonychus ulmi populations reached moderate levels only on Merlot and in the first part of experiments. The variety-phytoseiid species relationship was temporary as, at the end of experiments,T. pyri was completely dominant on all varieties. This new situation started when prey occurrence and interspecific competition decreased in importance. The moderate success of theT. pyri release on Merlot contrasts with the results of previous experiments. Two factors could be involved in this phenomenon: low interspecific competition by phytoseiids and predation by macropredators.Amblyseius aberrans was able to displaceA. andersoni andT. pyri on grape varieties where the two species were more abundant and reached higher population densities on varieties with pubescent leaf undersurfaces. In the first experimental year, spider mite densities were reduced more effectively inA. aberrans release plots than in the control or inT. pyri release plots. One year later,P. ulmi reached lower levels in the release treatments than in the control.Typhlodromus pyri andA. aberrans persisted in conditions of prey scarcity. The high competitivity ofA. aberrans over the remaining two phytoseiid species constitutes a major factor in selecting predatory species for inoculative releases in vineyards.     

Factors determining species displacement of related predatory mite species (Acari: Phytoseiidae)

Neoseiulus womersleyi (Acari: Phytoseiidae) used to be the dominant species in fruit-tree orchards throughout Japan, but starting in the 1990s, N. womersleyi began to be displaced by Neoseiulus californicus in central and southwestern Japan. The present study was conducted to examine factors explaining the displacement of N. womersleyi by N. californicus. First, we confirmed under laboratory conditions that N. californicus could exclude N. womersleyi if they initially coexisted in a 1:1 ratio. During a 2-h continuous observation period, none of the heterospecific pairs had copulated and after 5 days together with heterospecific males, none of the females had laid eggs. When these females were placed with conspecific males, normal numbers of offspring were produced. Moreover, conspecific matings were not substantially disturbed in the presence of heterospecific males or females. Total fecundity was significantly lower in N. womersleyi than in N. californicus, but their r _m values did not differ from each other. On the other hand, the frequency of intraguild predation by N. californicus on N. womersleyi was significantly higher than vice versa. From these results, we concluded that not reproductive interference nor differential female fecundity but asymmetrical intraguild predation seemed to explain the competitive exclusion of N. womersleyi by N. californicus.     

Efficacy of indigenous predatory mites (Acari: Phytoseiidae) against the citrus rust mite Phyllocoptruta oleivora (Acari: Eriophyidae): augmentation and conservation biological control in Israeli citrus orchards

The citrus rust mite (CRM), Phyllocoptruta oleivora (Acari: Eriophyidae) is a cosmopolitan key pest of citrus, inflicting severe economic damage if not controlled. In Israel, CRM damages all citrus cultivars. International regulation and increasing control failures of CRM led growers to seek sustainable biological control solutions such as acarine biological control agents. Laboratory studies conducted in Israel have indicated that the indigenous predator species Amblyseius swirskii, Iphiseius degenerans, Typhlodromus athiasae and Euseius scutalis (all Acari: Phytoseiidae) can potentially control CRM. Our general objective in the present study was to bridge the gap of knowledge between laboratory studies and the lack of control efficacy of these species in commercial orchards. Predator augmentation in the field showed that although predator populations increased immediately following releases they later decreased and did not affect CRM populations. When A. swirskii augmentation was combined with a series of maize pollen applications, A. swirskii populations were enhanced substantially and continuously but again CRM populations were not affected. Growth chamber studies with CRM-infested seedlings, with or without a maize pollen supplement, indicated that pollen provisioning led to population increase of E. scutalis and A. swirskii but only E. scutalis significantly lowered CRM populations. Control with E. scutalis was confirmed in the field on CRM infested seedlings with pollen provisioned by adjacent flowering Rhodes grass. While experiments in mature citrus orchard showed that pollen supplement usually increased predator populations they also indicated that other factors such as intraguild interactions and pesticide treatments should be taken into account when devising CRM biological control programs.     

Population dynamics of phytophagous and predatory mites (Acari: Tetranychidae, Eriophyidae, Phytoseiidae) on bamboo plants in Fujian, China

The seasonal cycle and population dynamics of Schizotetranychus nanjingensis Ma and Yuan, Aponychus corpuzae Rimando (Tetranychidae), Aculus bambusae Kuang (Eriophyidae) and their natural enemy Typhlodromus bambusae Ehara (Phytoseiidae) were studied during 1996–1998 in moso bamboo forests in Nanping, Fujian, China. Damage to bamboo leaves was often caused by mixed populations of the three phytophagous species, which displayed different seasonal dynamics: S. nanjingensis fed actively and reproduced in May and from late July to late October (aestivating from June to mid July), Ap. corpuzae was active and reproduced from mid July to late November, and Ac. bambusae from July to next February. During Spring S. nanjingensis was the dominant species with the greatest niche width, but in Summer Ac. bambusae and Ap. corpuzae became dominant and had greater niche widths. In Autumn, Ac. bambusae decreased but both S. nanjingensis and Ap. corpuzae mites increased and the latter became dominant. In winter all species decreased sharply in number. The two spider mite species (S. nanjingensis and Ap. corpuzae) had high niche overlaps in all four seasons. Niche overlaps between the eriophyid Ac. bambusae and the two spider mites were similarly high except during the spring when Ac. bambusae was absent. The predatory mite (T. bambusae) had higher niche overlap with Ap. corpuzae than with others during Autumn and Winter, but during Spring and Summer niche overlap was higher with S. nanjingensis.     

Biological studies of the Australian predatory mite Typhlodromalus lailae (Schicha) (Acari: Phytoseiidae)

Abstract The biology of the Australian phytoseiid mite Typhlodromalus lailae is described from material collected in Western Australia and New South Wales in 1994. At 25°C, when fed on cumbungi ( Typha sp.) pollen, the life cycle is completed in approximately 6 days, with an intrinsic rate of natural increase ( r _m ), of 0.38. Female−male pairs produced a mean total of 44.6 eggs within 22 days of oviposition, with 39% of these females living in excess of 29 days. Females that were deprived of males after first mating stopped laying eggs after 4−9 days, but if another male was added they resumed egg laying and produced close to a full complement of eggs (mean 42 eggs). At 25°C, fecundity on a diet of cumbungi pollen or thrips larvae (first-instar Frankliniella schutzei Trybom) was high, averaging 3.71 and 3.33 eggs per day, respectively, over a 3-day period. The sex ratio was approximately 1.5 females to 1 male. Consumption rate of thrips was also high, with an average of approximately seven first-instar or two second-instar F. schultzei larvae eaten per day. The species was also observed to feed on broad mite, Polyphagotarsonemus latus (Banks), and tomato russet mite, Aculops lycopersici (Massee). No diapause was observed under conditions of 25°C 8 h light and 10°C 16 h dark. Eggs were sensitive to low humidity, with 50% failing to hatch below 71.1% relative humidity. This species is of interest as a candidate biological control agent for thrips, broad mite and tomato russet mite in protected crops.     

Rapid cold hardening in the predatory mite Euseius (Amblyseius) finlandicus (Acari: Phytoseiidae)

A rapid cold hardening response was studied in diapause and non-diapause females of the predatory mite Euseius finlandicus. When laboratory reared diapause and non-diapause females were transferred and maintained from the rearing temperature of 20 degrees C for 2 h to -11.5 degrees C and -10 degrees C, 10 to 20% survived respectively. However, conditioning of diapause females for 4 h at a range of temperatures from 0 to 10 degrees C before their exposure for 2 h to -11.5 degrees C, increased survival to approximately 90%. Similarly, conditioning of non-diapause females for 4 h at 5 degrees C before their exposure for 2 h to -10 degrees C increased survival to 90%. A similar rapid cold hardening response in both diapause and non-diapause females was also induced through gradual cooling of the mites, at a rate of approximately 0.4 degrees C per min. The rapid increase in cold tolerance after prior conditioning of the mites to low temperatures, was rapidly lost when they returned to a higher temperature of 20 degrees C. Rapid cold hardening extended the survival time of diapause and non-diapause females at sub-zero temperatures. The cost of rapid cold hardening in reproductive potential after diapause termination was negligible. In non-diapause females, however, the increase in cold tolerance gained through gradual cooling could not prevent cold shock injuries, as both fecundity and survival were reduced.     

Distortion of the spermathecae of phytoseiid mites (Acari: Phytoseiidae) in slide preparation

The shape and dimensions of the spermathecae are taxonomically important characteristics in phytoseiid mites. In experiments with Neoseiulus californicus (McGregor) and Typhlodromus pyri Scheuten, the shape of the cervix of the spermatheca changed considerably and its diameter was increased by 42–49% when mites were flattened, compared to being mounted with 140m spacers underneath the coverslips. Dimensions of the dorsal shield were also affected by flattening, increasing by 3–5%.     

Susceptibility to deltamethrin in the predatory mites Neoseiulus californicus and Phytoseiulus macropilis (Acari: Phytoseiidae) populations in protected ornamental crops in Brazil

Knowledge of inter and intra-specific variation in the susceptibility of natural enemies to pesticides could help to better design integrated pest management strategies. The objective of this research was to evaluate the susceptibility to deltamethrin in populations of the predatory mites Neoseiulus californicus (McGregor) and Phytoseiulus macropilis (Banks) populations collected from protected ornamental crops in Brazil. The susceptibility to deltamethrin was characterized against immature and adult stages of both species. The impact of this insecticide was also measured by estimating the intrinsic rate of increase (r _i). The immature and adult stages of N. californicus were approximately 3,600 and 3,000-fold more tolerant to deltamethrin than those of P. macropilis. However, high variability in the susceptibility to this insecticide was detected among P. macropilis populations, with resistance ratios of up to 3,500-fold. The selection of deltamethrin-resistant strains of P. macropilis could be exploited in applied biological control programs.