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.     

Within-Plant Migration of the Predatory Mite Typhlodromalus aripo from the Apex to the Leaves of Cassava: Response to Day–Night Cycle, Prey Location and Prey Density

Under attack by herbivores, plants produce a blend of “herbivore-induced plant volatiles (HIPV)” that help natural enemies of herbivores locating their prey, thereby helping plants to reduce damage from herbivory. The amount of HIPV emitted by plants increases with herbivore density and is positively correlated with the intensity of the olfactory response of natural enemies. In this study, we determined the effects of density or within-plant distribution of the herbivorous mite Mononychellus tanajoa on movement of the predatory mite Typhlodromalus aripo out of apices of cassava plants. Proportions of T. aripo that migrated out of apex, and distances traveled were significantly higher when M. tanajoa was further away from the apex—i.e. on middle or bottom leaves of cassava plants—than when present on top leaves, or absent from the plant. This supports previous field observations that T. aripo is not a sit-and-wait predator but uses HIPV to search and locate its prey within cassava plant.     

Attraction of the predatory mites Typhlodromalus manihoti and Typhlodromalus aripo to cassava plants infested by cassava green mite

The attraction of the predatory mites, Typhlodromalus manihoti and Typhlodromalus aripo, to the host plant-spider mite complex, Manihot esculenta – Mononychellus tanajoa, was investigated with a Y-tube olfactometer. Factors examined included predator starvation period, several combinations of cassava leaf biomass and initial M. tanajoa infestations, M. tanajoa-damaged leaves with mites and/or their residues removed, M. tanajoa alone, and mechanically damaged cassava leaves. We found that females of T. manihoti and T. aripo were significantly attracted to M. tanajoa-infested cassava leaves when the predators were starved for 2, 6, or 10 h. Satiated T. aripo was significantly attracted to infested cassava leaves whereas satiated T. manihoti did not discriminate between infested and non-infested leaves. When a choice was given between either two or four leaves infested with 200 female M. tanajoa and an equivalent number of non-infested leaves, 2 h-starved T. manihoti and T. aripo were significantly attracted to each of the infested groups of cassava leaves. At a density of 12 female M. tanajoa per leaf on four leaves, 2 h-starved T. manihoti was still attracted to M. tanajoa-infested leaves whereas 2 h-starved T. aripo was not attracted. When a choice was given between non-infested cassava leaves and either infested leaves from which only M. tanajoa females had been removed, or infested leaves from which all M. tanajoa and their visible products (web, feces) had been wiped off, T. aripo preferred odors from both types of previously infested leaves. Typhlodromalus manihoti was only attracted to infested leaves from which the M. tanajoa females only had been removed. Finally, the two predators were not attracted to 400 female M. tanajoa on clean cotton wool or to mechanically wounded leaves. This supports the hypothesis that M. tanajoa damage induces volatile cues in cassava leaves that attract T. manihoti and T. aripo to M. tanajoa-infested leaves.     

Plant feeding by a predatory mite inhabiting cassava

Plant feeding by arthropod predators may strongly affect the dynamics of bi- and tri-trophic interactions. We tested whether a predatory mite, Typhlodromalus aripo, feeds upon its host plant, cassava. This predator species is an effective biological control agent of Monoychellus tanajoa (the cassava green mite or CGM) a herbivorous mite specific to cassava. We developed a technique to detect plant feeding, based on the use of a systemic insecticide. We found that T. aripo feeds upon plant-borne material, while other predatory mite species, Neoseiulus idaeus and Phytoseiulus persimilis, do not. Subsequently, we measured survival of juveniles and adult females of T. aripo and N. idaeus, both cassava-inhabiting predator species, on cassava leaf discs. Survival of T. aripo was higher than that of N. idaeus. Thus, T. aripo was able to withstand longer periods of prey scarcity. Because CGM populations fluctuate yearly and are heterogeneously distributed within plants, plant feeding may facilitate the persistence of populations of T. aripo in cassava fields and its control of CGM outbreaks. This revised version was published online in July 2006 with corrections to the Cover Date.     

Interactions Between Two Neotropical Phytoseiid Predators on Cassava Plants and Consequences for Biological Control of a Shared Spider Mite Prey: a Screenhouse Evaluation

The issue of introducing single or multiple natural enemy species for classical biological control has been an area of intense inquiry by ecologists and biological control practitioners. This is particularly relevant to classical biological control of cassava green mite Mononychellus tanajoa (Bondar) (Tetranychidae) in Africa, as this pest mite is shared by several natural enemies in the Neotropics (its area of origin), two of which have been introduced and established widely in Africa. We conducted two screenhouse experiments using the two neotropical phytoseiid predatory mites, Typhlodromalus aripo DeLeon and Typhlodromalus manihoti Moraes, to determine the effects of single and two-predator species on population dynamics of the two predators and on suppression of M. tanajoa populations. The two predators are thought to be complementary in their impact on their shared prey M. tanajoa, due to similarities in their preference for this prey and to differences in their spatial distribution and foraging activities on cassava. The two predator species were released alone or together at low and at high initial densities of M. tanajoa. In all cases, predator releases resulted in significant suppression of M. tanajoa, but the degree of suppression did not differ among single and two-species releases with one exception: at high initial density of M. tanajoa, releases of T. aripo alone had less impact than that of either T. manihoti alone or of the two species together. Typhlodromalus aripo also appeared to be inferior as a competitor of T. manihoti: at low initial density of M. tanajoa, the proportion of T. aripo in the two-predator release treatments gradually declined and was strikingly lower than in the single species release, probably due to intraguild predation on its larvae by T. manihoti. However, T. aripo persisted longer than T. manihoti after elimination of M. tanajoa. On the basis of this study under semi-natural conditions, it appears that either species is sufficient for controlling M. tanajoa populations, with T. manihoti being more efficient at high initial prey densities and T. aripo at low initial prey densities. At high prey density, T. manihoti increased to large numbers and outcompeted T. aripo. Relevance of these findings to larger spatial scale and under natural conditions is discussed.     

Single versus multiple enemies and the impact on biological control of spider mites in cassava fields in West-Africa

To determine whether to use single or multiple predator species for biological pest control requires manipulative field experiments. We performed such tests in Benin (West Africa) in cassava fields infested by the cassava green mite Mononychellus tanajoa, and the cotton red mite Oligonychus gossypii. These fields also harboured the cassava apex-inhabiting predator Typhlodromalus aripo and either the leaf-inhabiting predator Amblydromalus manihoti or Euseius fustis. We manipulated predator species composition on individual plants to determine their effect on prey and predator densities. In fields with T. aripo plus A. manihoti, M. tanajoa densities were reduced by T. aripo alone or together with A. manihoti, but neither of these predators, alone or together, reduced O. gossypii densities. In fields with T. aripo plus E. fustis, T. aripo alone or together with E. fustis exerted significant control over O. gossypii, but weak control over M. tanajoa. Densities of any of the predator species were not affected by co-occurring predator species, suggesting a minor role for intraguild predation in the field, contrary to earlier experiments on small plants in the laboratory. We conclude that (1) T. aripo is the most effective predator species in suppressing M. tanajoa, (2) two predator species, T. aripo and E. fustis, are needed to suppress O. gossypii, and (3) predator species together on the same plant do not negatively affect each other nor the extent to which they control their prey. We argue that intraguild predation is reduced due to partial niche separation among predator species.     

Mix‐planting pubescent and glabrous cassava affects abundance of Typhlodromalus aripo and its prey mite Mononychellus tanajoa

There is an increasing awareness that vegetation diversity can affect herbivore and natural enemy abundance and that plants can play a major role in directly manipulating natural enemy abundance for protection against herbivore attacks. Using data from cassava fields, we aimed at (i) testing the capacity of the predatory mite Typhlodromalus aripo to control the herbivorous mite Mononychellus tanajoa in a chemical exclusion trial; and (ii) testing, based on the differential preference by T. aripo for cassava cultivars, how combinations of two morphologically different cassava cultivars with differential suitability to the predator can improve its population densities on the non‐favourable cultivar, thereby reducing M. tanajoa densities with subsequent increases in cassava yield. The study was conducted in a cassava field in Benin, West Africa. The experiments confirmed that T. aripo effectively suppresses M. tanajoa populations on both cultivars and showed, in the no‐predator‐exclusion experiments, that cultivar combinations have significant effects on M. tanajoa and T. aripo densities. Indeed, T. aripo load on the non‐preferred cultivar was lowest in subplots where the proportion of T. aripo‐preferred cultivar was also low, while, and as expected, M. tanajoa load on the non‐preferred cultivar showed decreasing trends with increasing T. aripo densities. The possible mechanisms by which cultivar mixing could increase predator load on the non‐favourable cultivar were discussed. Our data showed that appropriate cultivar combinations effectively compensate for morphologically related differences in natural enemy abundance on a normally predator‐deficient cultivar, resulting in lower pest densities on the non‐favourable cultivar. In practical terms, this strategy could, in part, enhance adoption of cultivars that do not support sufficient levels of natural enemies for pest control.     

Living at the threshold: Where does the neotropical phytoseiid mite Typhlodromalus aripo survive the dry season?

The establishment of the neotropical predatory mite Typhlodromalus aripo in sub-Saharan Africa has resulted in broadly successful biological control of the cassava green mite Mononychellus tanajoa throughout the cassava belt of Africa. In some mid-altitude areas and drier lowland savannahs of sub-Saharan Africa, which are characterized by cool or hot long (≥5 months) dry seasons, the predator disappears from its habitat in the cassava apex during the dry season and reappears after the onset of rains. It is not known, however, where the predator remains during this time period. In this study, we conducted a field enclosure experiment of cassava plants with the objectives to determine if (a) T. aripo survives at very low densities in the apex, if (b) it survives in the soil or leaf litter below the cassava plant, and if (c) it recolonizes the cassava plant from the surrounding vegetation. Towards the end of the dry season, when the predators had disappeared from all cassava plants included in the experiment, five treatments were applied: (1) plants without enclosure; (2) plants with enclosure; (3) plants with enclosure, apices removed; (4) plants with enclosure, glue barrier around stem; and (5) plants kept free of T. aripo, without enclosure. Predator (re)appearance on cassava apices was monitored non-destructively at weekly intervals and was expressed as the proportion of plants with at least one apex with T. aripo per total number of plants of the treatment. The predators reappeared first on the plants of the treatments (1), (2), and (4). With a time lag of 7–8 weeks, the predators appeared also on the plants of the treatments (3) and (5). The time pattern of the predator’s (re)appearance in the cassava apex of the different treatments suggests that (a) T. aripo survives the dry season in very low densities in the cassava apex; this result is supported by an assessment of the efficiency of non-destructive visual in-field apex inspections which proved that about 10% of the cassava apices that had T. aripo were not recognized as such; (b) T. aripo does not survive in the soil or leaf litter, but we did document cases in a screenhouse experiment, where few individuals migrated down to the ground and walked over exposed soil until they reached the apex bouquet traps; additionally, microclimate measurements in various cassava plant strata proved that the cassava apex and the cassava stem base are the locations with the highest relative humidity during the dry season—which makes the stem base a potentially interesting refuge; (c) T. aripo does not survive in the surrounding vegetation, which is supported by a vegetation survey, where T. aripo was not found on any other plant species than cassava.     

Cannibalism and Interspecific Predation in a Phytoseiid Predator Guild from Cassava Fields in Africa: Evidence from the Laboratory

Interspecific predation and cannibalism are common types of interaction in phytoseiid predator guilds, but the extent and nature of these interactions have not been determined yet in phytoseiid guilds composed of African native and neotropical exotic phytoseiid predators found in cassava habitat in southern Africa. We determined in laboratory experiments the level of cannibalism and interspecific predation among the three phytoseiid mite species Euseius fustis, Iphiseius degenerans, and Typhlodromalus aripo in the absence of food and in the presence of limited or abundant quantities of two food types – Mononychellus tanajoa and maize pollen – commonly found on cassava in Africa. When confined without food, only two T. aripo females laid each two eggs within 5 days, and this species survived longer than I. degenerans and E. fustis. In the presence of con- or hetero-specific larvae or protonymphs, the three species fed more on the former than on the latter, and more on hetero-specifics than on con-specifics. Oviposition rates of the three species did not exceed 0.7 egg/female/day on con- and hetero-specific immatures. Typhlodromalus aripo and E. fustis survived longer on con-specific and hetero-specific larvae and on hetero-specific protonymphs than in the absence of any food, while T. aripo survived longer than the two other species on the same diets. Provision of limited quantity of food decreased interspecific predation rate by I. degenerans and T. aripo, but not by E. fustis, and increased oviposition rate and longevity of all three species. Provision of abundant food, however, eliminated cannibalism by all three species and further reduced interspecific predation rates, but their oviposition and longevity remained relatively unchanged compared with limited food provision. Potential consequences of cannibalism and interspecific predation among phytoseiid mites on cassava for the biological control of M. tanajoa are discussed.     

The predatory mite Typhlodromalus aripo prefers green-mite induced plant odours from pubescent cassava varieties

It is well known that plant-inhabiting predators use herbivore-induced plant volatiles to locate herbivores being their prey. Much less known, however, is the phenomenon that genotypes of the same host plant species vary in the attractiveness of these induced chemical signals, whereas they also differ in characteristics that affect the predator’s foraging success, such as leaf pubescence. In a series of two-choice experiments (using a Y-tube olfactometer) we determined the preference of Typhlodromalus aripo for pubescent versus glabrous cassava cultivars infested with the cassava green mite Mononychellus tanajoa and also the preference for cultivars within each of the two groups. We found that when offered a choice between pubescent and glabrous cassava cultivars (either apex or leaves), T. aripo was significantly more attracted to pubescent cultivars. For each cultivar, M. tanajoa infested leaves and apices were equally attractive to T. aripo. There was however some variation in the response of T. aripo to M. tanajoa-infested plant parts within the group of pubescent cultivars, as well as within the group of glabrous cultivars. Our study confirms not only that T. aripo uses herbivore-induced plant volatiles to search for prey in cassava fields, but it also shows that it can discriminate between glabrous and pubescent cultivars and prefers the latter. This knowledge can be useful in selecting cultivars that are attractive and suitable to T. aripo, which, in turn, may promote biological control of the cassava green mite.     

Effects of prey mite species on life history of the phytoseiid predators Typhlodromalus manihoti and Typhlodromalus aripo

The effects of prey mite suitability on several demographic characteristics of phytoseiid predators and the relationship of these effects to the potential of phytoseiid predators to control herbivorous mite populations are well documented. Evidence has also accumulated in the last 20 years demonstrating that phytoseiid predators utilize herbivorous prey mite-induced plant volatiles as olfactory cues in locating their herbivorous mite prey, but less well established is the predictability of reproductive success from the ability of the predators to utilize olfactory cues to locate their prey, and how these processes are related to the success of the predators as biological control agents of the herbivorous mite. In this study, we determined in laboratory no choice experiments, the development, survivorship and fecundity of the two neotropical phytoseiid predators Typhlodromalus manihoti Moraes and T. aripo DeLeon when feeding on three herbivorous mites, including the key prey species Mononychellus tanajoa (Bondar), and the two alternative prey species Oligonychus gossypii (Zacher) and Tetranychus urticae (Koch). Intrinsic rate of increase (r_m) of T. aripo was 2.1 fold higher on M. tanajoa as prey compared with T. urticae as prey, while it was almost nil on O. gossypii. For T. manihoti, r_m was 2.3 fold higher on M. tanajoa as prey compared with O. gossypii as prey, while reproduction was nil on T. urticae. An independent experiment on odor-related prey preference of the two predator species showed that T. manihoti and T. aripo preferred odors from M. tanajoa-infested leaves to odors from O. gossypii-infested leaves. Moreover, both predator species preferred odors from M. tanajoa-infested leaves over those from T. urticae-infested leaves. As reported here, life history of the two predatory mites matches odor-related prey preference if the key prey species is compared to the two inferior prey species. The implications of our findings for the persistence of T. manihoti and T. aripo and biological control of M. tanajoa in the cassava agroecosystem in Africa are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of the entomopathogenic fungus Neozygites tanajoae and the predatory mite Typhlodromalus aripo on cassava green mite densities: screenhouse experiments

To improve biocontrol of the cassava green mite, Mononychellus tanajoa, a series of screenhouse experiments were conducted on cassava to determine the effects of single and combined releases of the predatory mite Typhlodromalus aripo and the entomopathogenic fungus Neozygites tanajoae on the suppression of M. tanajoa. We showed that, separately or in combination, T. aripo and N. tanajoae significantly reduced M. tanajoa populations. Moreover, combined release of T. aripo and N. tanajoae on the same cassava plants improved M. tanajoa control. However, our data suggest between T. aripo and N. tanajoae an asymmetric competition that significantly affected N. tanajoae only. This interference is indirect, probably mediated by their common prey or host mite, M. tanajoa since N. tanajoae is not pathogenic to T. aripo. We conclude that adding N. tanajoae to T. aripo populations in Africa would not negatively affect T. aripo and would further promote biocontrol of M. tanajoa.     

Infochemical-Mediated Niche Use by the Predatory Mites Typhlodromalus manihoti and T. aripo (Acari: Phytoseiidae)

In Africa, Typhlodromalus manihoti and T. aripo, two introduced predators of the cassava green mite Mononychellus tanajoa, occupy different parts of cassava foliage. In the present study, niche use by these two predators, as mediated by prey-induced infochemicals, was investigated. In response to prey feeding damage, cassava plant parts emit volatile blends, that attract phytoseiidae predators. When given a choice between old cassava leaves infested with M. tanajoa and either apices or young cassava leaves infested with M. tanajoa, T. aripo displayed a marked preference for odors emitted from either infested apices or infested young leaves over infested old leaves but showed no preference for odors from apices versus young leaves, all infested with M. tanajoa. Typhlodromalus manihoti did not discriminate between volatiles from the three infested cassava plant parts. Our data show that T. aripo uses differences in volatile blends released by infested cassava plant parts and restricts its fundamental niche to a realized niche, which enables coexistence with its competitor T. manihoti.     

Effect of some food sources associated with cassava in Africa on the development, fecundity and longevity of Euseius fustis (Pritchard and Baker) (Acari: Phytoseiidae)

Various foods associated with cassava were tested for their effect on the development, fecundity and longevity of Euseius fustis, the most common phytoseiid species found on cassava in Africa. Euseius fustis developed successfully to adulthood on the spider mite prey species Mononychellus tanajoa (Bondar) and Oligonychus gossypii (Zacher) and on pollen from maize, castor bean and cassava. Euseius fustis also completed development on water-diluted phloem exudate from cassava, diluted honeydew from the cassava mealybug and on various pollen and prey combinations. When reared on Tetranychus urticae Koch prey or free water only, E. fustis did not develop past the deutonymphal stage. All larvae held on clean leaf discs on water-soaked cotton died without moulting, suggesting that E. fustis must feed in order to moult to the nymphal stages. Diets of maize plus castor bean pollen and maize pollen plus M. tanajoa resulted in the highest rate of development, the highest fecundity and the greatest longevity. Castor bean pollen alone and maize pollen alone produced a higher fecundity and greater longevity than M. tanajoa tested alone. A colony of E. fustis reared continuously for seven generations on castor bean pollen produced nine times more adult females than a colony of E. fustis reared continuously on M. tanajoa. No negative effects on the development and fecundity of E. fustis were observed after seven generations were reared on pollen.     

Functional responses of Iphiseius degenerans and Neoseiulus teke (Acari: Phytoseiidae) to changes in the density of the cassava green mite, Mononychellus tanajoa (Acari: Tetranychidae)

The functional responses of protonymph and adult female Iphiseius degenerans and Neoseiulus teke to increasing density of three stages of their prey, the cassava green mite (CGM), Mononychellus tanajoa, were studied on excised cassava leaf discs under laboratory conditions. The responses obtained were predominantly sigmoid type III curves with the highest plateau when both stages of I. degenerans and N. teke were preying on CGM eggs. In all cases, the predation rate of the former species exceeded that of the latter. The empirical data were fitted by four different models. From the models, the attack coefficient (a) and handling time (T _h) were estimated. For a given predator stage (protonymph or adult female), the predator's attack coefficient declines and handling time increases as the prey gets larger. For a given prey stage, the predator's attack coefficient increases and handling time decreases as the predator stage becomes larger.     

Reproductive responses of Iphiseius degenerans and Neoseiulus teke (Acari: Phytoseiidae) to changes in the density of the cassava green mite, Mononychellus tanajoa (Acari: Tetranychidae)

We assessed the reproductive responses of adult female Iphiseius degenerans and Neoseiulus teke to increasing density of three stages of their prey, Mononychellus tanajoa, on cassava leaf discs under laboratory conditions. The oviposition rates increased with number of prey consumed per predator per day with a maximum of approximately two eggs per day for I. degenerans and four eggs per day for N. teke. The oviposition rate of N. teke was higher when consuming eggs than other prey stages. Neoseiulus teke was more efficient than I. degenerans in converting consumed prey into egg production. The data were adequately described by simple mathematical models.     

Flexible antipredator behaviour in herbivorous mites through vertical migration in a plant

When predation risk varies in space and time and with predator species, successful prey defence requires specific responses to each predator. In cassava fields in Africa, the herbivorous cassava green mite (Mononychellus tanajoa) is attacked by three predatory mite species that are segregated within the plant: the leaf-dwelling Typhlodromalus manihoti and Euseius fustis occur on the middle leaves, whereas the apex-inhabiting T. aripo migrates from the apex to the top leaves only during the night. We found that differential distributions of these predators allow prey to escape predation by vertical migration to other plant strata. We studied the role of odours in the underlying prey behaviour on predator-free plants placed downwind from plants with predators and prey or with prey only. Prey showed increased vertical migration in response to predator-related odours. Moreover, these responses were specific: when exposed to odours associated with T. manihoti, prey migrated upwards, irrespective of the plant stratum where they were placed. Odours associated with T. aripo triggered a flexible response: prey on the top leaves migrated downwards, whereas prey on the middle leaves migrated upwards. Odours associated with E. fustis, a low-risk predator, did not elicit vertical migration. Further experiments revealed that: (1) prey migrate up or down depending on the stratum where they are located, and (2) prey discrimination among predators is based upon the perception of predator species-specific body odours. Thus, at the scale of a single plant, odour-based enemy specification allows herbivorous mites to escape predation by vertical migration.     

Effect of cassava exudate and prey densities on the survival and reproduction of Typhlodromalus limonicus (Garman & McGregor) s.l. (Acari: Phytoseiidae), a predator of the cassava green mite,Mononychellus tanajoa (Bondar) (Acari: Tetranychidae)

The effects of cassava exudate and prey densities on reproduction and survival of the predatory mite, Typhlodromalus limonicus (Garman & McGregor) (Acari: Phytoseiidae), were investigated in the laboratory. Females were provided either cassava exudate ad lib. daily, low or high numbers of the cassava green mite prey, Mononychellus tanajoa (Bondar) (Acari: Tetranychidae) daily, or exudate for 5 or 10 days before switching to a low or high prey diet. Females fed only exudate laid no eggs. Females fed exudate before prey experienced a significant decrease (30%) in the number of eggs laid compared to females fed high numbers of prey daily. The reduction in fecundity was the result of prolonged preoviposition periods (2.0 days on prey daily vs 4.0 days on exudate before prey) and reduced number of eggs laid per female per day (1.7 eggs per female per day on prey daily vs 0.4 eggs per female per day on exudate before prey). Females fed only exudate had a greater survival rate and longevity than females fed prey daily or females fed exudate before a diet of prey. These results suggest that T. limonicus can survice for a limited period on cassava exudate during periods of low prey availability, but requires prey to complete oögenesis and propagate the population.     

Relative contribution of biotic and abiotic factors to the population density of the cassava green mite, Mononychellus tanajoa (Acari: Tetranychidae)

The cassava green mite, Mononychellus tanajoa, is a key pest of cassava, Manihot esculenta Crantz (Euphorbiaceae), and it may be kept in check by naturally occurring predatory mites of the family Phytoseiidae. In addition to predatory mites, abiotic factors may also contribute to regulate pest mite populations in the field. Here, we evaluated the population densities of both M. tanajoa and the generalist predatory mite Euseius ho DeLeon (Acari: Phytoseiidae) over the cultivation cycle (11 months) of cassava in four study sites located around the city of Miranda do Norte, Maranhão, Brazil. The abiotic variables rainfall, temperature and relative humidity were also recorded throughout the cultivation cycle of cassava. We determined the relative importance of biotic (density of E. ho) and abiotic (rainfall, temperature and relative humidity) factors to the density of M. tanajoa. The density of M. tanajoa increased whereas the density of E. ho remained constant throughout time. A hierarchical partitioning analysis revealed that most of the variance for the density of M. tanajoa was explained by rainfall and relative humidity followed by E. ho density and temperature. We conclude that abiotic factors, especially rainfall, were the main mechanisms driving M. tanajoa densities.     

Functional response of Euseius concordis to densities of different developmental stages of the cassava green mite

Both prey density and developmental stage of pests and natural enemies are known to influence the effectiveness of biological control. However, little is known about the interaction between prey density and population structure on predation and fecundity of generalist predatory mites. Here, we evaluated the functional response (number of prey eaten by predator in relation to prey density) of adult females and nymphs of the generalist predatory mite Euseius concordis to densities of different developmental stages of the cassava green mite Mononychellus tanajoa, as well as the fecundity of adult females of the predator. We further assessed the instantaneous rate of increase, based on fecundity and mortality, of E. concordis fed on eggs, immatures and adults of M. tanajoa. Overall, nymphs and adults of E. concordis feeding on eggs, immatures and females of M. tanajoa had a type III functional response curve suggesting that the predator increased prey consumption rate as prey density increased. Both nymphs and adult females of the predator consumed more eggs than immatures of M. tanajoa from the density of 20 items per leaf disc onwards, revealing an interaction between prey density and developmental stage in the predatory activity of E. concordis. In addition, population growth rate was higher when the predator fed on eggs and immatures in comparison with females. Altogether our results suggest that E. concordis may be a good candidate for the biological control of M. tanajoa populations. However, the efficiency of E. concordis as a biological control agent of M. tanajoa is contingent on prey density and population structure.