Pathogenicity and specificity of Neozygites tanajoae and Neozygites floridana (Zygomycetes: Entomophthorales) isolates pathogenic to the cassava green mite

The cassava green mite (CGM), Mononychellus tanajoa, a native of South America was accidentally introduced into Africa where it causes serious crop losses. The possibility of introducing classical biological agents from the native home of CGM into Africa was investigated. Thus, we conducted a series of laboratory assays of the native fungal pathogens, Neozygites tanajoae from Brazil and Neozygites floridana from Colombia and Brazil, and compared them with N. tanajoae isolates from Benin. Infectivity of both fungal species, was assayed against the twospotted spider mite, Tetranychus urticae, and against the red mite, Oligonychus gossypii. Pathogenicity against CGM and host range studies were conducted by transferring adult females of each mite species to leaf discs containing sporulated cadavers with a halo of conidia of each fungal isolate. All isolates caused some degree of infectivity to CGM. None of the isolates of N. floridana and N. tanajoae tested were pathogenic to O. gossypii, and only two isolates infected T. urticae. Most isolates from Brazil were highly virulent and infected only CGM. Sixteen N. tanajoae isolates caused more than 89% mortality and more than 62% of the CGM became mummified. A mummified CGM is characteristically a swollen, brown fungus-killed mite that has great potential to produce conidia. However, high mortality was not always associated with high mummification. The median mummification time ranged from 4.4 to 6.7 days. Five Brazilian isolates caused >75% mummification with a median mummification time <5 days. Isolates that cause high mummification in a short period of time would be more likely to cause epizootics and to establish in the new environment. Therefore, these isolates would be the best candidates for introduction to Africa.     

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.     

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.     

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.     

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.     

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.     

Age-dependent rates of infection of cassava green mites by a fungal pathogen in Brazil

Age-specific effects of invertebrate pathogens on their hosts can greatly influence the population dynamics in such interactions. Explanations for such differences are usually sought within differing intrinsic susceptibilities of the host life stages but we present data which indicate that host size, behaviour and life history may be the overriding factors determining age-specific effects of a fungal pathogen, Neozygites floridana (Entomophthorales: Neozygitaceae) on spider mites (Mononychellus tanajoa Bondar, Acari: Tetranychidae). Epizootics of N. floridana in spider mites are characterised by much greater relative mortality of adult females compared with other life stages (ca. 99%), despite similar physiological susceptibilities. We present empirical data that demonstrate encounter rates of mites with N. floridana increasing with life stage during an epizootic on cassava in northeastern Brazil. Estimates of the size, walking speeds and patterns, and life history of different life stages (and adult sexes) were used to calculate expected relative encounter rates which were found not to be different from the observed values (although not testable for larvae). This helps explain the different apparent susceptibility of host life stages in the field. Given the low ecological susceptibility of younger life stages to this pathogen, we predict that the interaction time between host and pathogen, determined by climatic conditions, will be critical in determining the degree of host population control in an epizootic. We further hypothesise that such variation in ecological susceptibility to pathogens can generate selection pressures on basic host traits, contributing to the sessile nature of many microarthropods. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Effect of temperature,humidity and photoperiod on mortality of Mononychellus tanajoa (Acari: Tetranychidae) infected by Neozygites cf. floridana (Zygomycetes: Entomophthorales)

The effect of temperature, humidity and photoperiod on the development of Neozygites cf. floridana (Weiser and Muma) in the cassava green mite, Mononychellus tanajoa (Bondar) was studied in the laboratory. Dead infected mites began to appear 2.5 days after inoculation. At 33 and 28°C peak mortalities were higher and occurred earlier (after 2.5 days), than at 23 and 18°C. Mean LT₅₀ (time for half the infected mites to die) decreased with increasing temperature as follows: 3.9, 3.0, 2.9 and 2.5 days at 18, 23, 28 and 33°C, respectively. When placed under conditions of high relative humidity for a period of 24 h, the percentage of dead infected mites from which the fungus sporulated was highest at 28°C (51.4%) and lowest at 33°C (6.5%). The development of the fungus inside the mite was not significantly affected by ambient humidity or photoperiod. No significant interactions between tested factors were found.     

Interactions of two natural enemies of Tetranychus evansi, the fungal pathogen Neozygites floridana (Zygomycetes: Entomophthorales) and the predatory mite, Phytoseiulus longipes (Acari: Phytoseiidae)

Tetranychus evansi is an exotic pest of Solanaceous crops in Africa discovered in Zimbabwe in 1979. Two natural enemies, the predatory mite Phytoseiulus longipes and the fungal pathogen Neozygites floridana are important causes of mortality in T. evansi populations in Brazil. The first part of this study assessed the effects of N. floridana on predation and oviposition of P. longipes fed on N. floridana infected T. evansi and T. urticae. No N. floridana hyphal bodies were found in P. longipes after this feeding, demonstrating that N. floridana is not pathogenic to P. longipes and does not affect its oviposition. The second part of the study investigated the time spent on searching for and consuming of eggs on leaf discs with and without N. floridana capilliconidia. Both the searching and the feeding time on the first egg were similar on leaf discs with and without capilliconidia. When P. longipes was offered the choice of feeding on eggs on leaf discs with or without capilliconidia, the numbers of eggs consumed were not different. The only N. floridana effect observed on P. longipes was reduced egg predation. In addition, increased time spent grooming on leaf discs with capilliconidia was observed. P. longipes was efficient in removing most capilliconidia attached to the body through self-grooming behavior. This suggests that although the predator did not avoid areas with capilliconidia, it detected and removed most capilliconidia attached to the body. Increased grooming may account for the lower egg predation rates.     

Seasonal cycles and persistence in an acarine predator-prey system on cassava in Africa

We applied time series analysis and a mechanistic predator-prey model to long-term data of monthly population counts of the herbivorous pest mite Mononychellus tanajoa and its introduced phytoseiid predator Typhlodromalus aripo from a cassava field in Benin, West Africa. In this approach, we determined the extent to which the main features of the observed predator-prey fluctuations in cassava fields can be explained from biotic traits inherent to the biology of predator and prey, and the extent of the significance of abiotic factors in determining population levels. The time series analyses with cross-correlation showed that the period of predator-prey fluctuations coincided with the annual pattern of intense rainfall and onset of dry season. A pronounced M. tanajoa peak followed after a short lag (2 weeks) by a T. aripo peak coincided with a trough in rainfall intensity. Both the prey and predator had local and lower peaks that coincided with high rainfall intensity, but with a considerably longer lag (ca. 3 months) compared with the high peaks occurring at the onset of the dry season. Regression of log-transformed data series (over a 7-year period) showed that—except for the first year after predator release—M. tanajoa fluctuated around an almost time-invariant mean population density, while T. aripo densities showed a consistent decline over the full observation period. To explain observed trends and periodic components in the data-series of predator and prey densities, we review hypotheses that are based on (1) the annual patterns and trends in abiotic factors, (2) mechanisms endogenous to the predator-prey system and (3) a combination of exogenous and endogenous factors.     

Survival of Neozygites cf. floridana (Zygomycetes: Entomophthorales) in mummified cassava green mites and the viability of its primary conidia

The survival of Neozygites cf. floridana (Weiser and Muma) as dry hyphal bodies in mummified cassava green mites, Mononychellus tanajoa (Bondar), at 5.0% RH in the dark was affected by storage temperature. Survival of the fungus in mummies kept at 24±1.0°C could be demonstrated for 6–7 months. When stored at 4°C, the fungus sporulated from 90% of the mummies liberating an average of 186.9 primary conidia per mummy even after a storage period of 16 months, when the experiment was terminated. The temperature, humidity and light condition significantly affected the viability of primary conidia. The percent viability across all factors dropped from 98.4% after 0 h (beginning of the experiment) to 23.4% after a 1 h exposure to the conditions tested. Lower temperatures maintained higher viabilities with 86.3% of the conidia surviving after 18 h at 18°C, whereas almost all conidia died after 12 h at 33°C. Conidia survived less than 1 h when exposed to SDs (saturation deficit) of 2.0 mm Hg or higher at any tested temperature.     

Interactive effects of selected pesticides on the two-spotted spider mite and its fungal pathogen Neozygites floridana

Benomyl affected populations of Tetranychus urticae by interfering with the pathogenic fungus, Neozygites floridana. Benomyl delayed but prolonged spider mite outbreaks. Few mites were infected with the pathogen when benomyl was used. Reductions in mite populations treated with fentin hydroxide were associated with a high incidence of N. floridana infection. Benomyl did not affect sporulation of N. floridana but appeared to inhibit conidial germination or growth of the fungus.

---

Résumé Le bénomyl a modifié les populations de Tetranychus urticae Koch en interférant avec son champignon pathogène, Neozygites floridana (Weiser & Muma). Le bénomyl retardait mais prolongeait les pullulations de l'acarien. Peu d'acariens étaient infectés par le champignon quand on utilisait du bénomyl. Les réductions des populations d'acariens traitées avec l'hydroxyde de fentine étaient associées à un haut niveau d'infection par N. floridana. Le bénomyl ne modifiait pas la sporulation de N. floridana mais semblait inhiber la germination des conidies ou la croissance du champignon.

     

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.     

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.     

Efficacy of the Ventral Abdominal Secretion of the Cockroach Eurycotis floridana (Blattaria: Blattidae) as a Defense Allomone

In assays investigating the abdominal sternal secretion of the cockroach Eurycotis floridana, it was determined that the secretion provides an effective deterrent against potential predators. Analysis of grooming patterns of Peromyscus leucopus mice revealed significant increases in mouth grooming and loss of coordination following exposure to E. floridana secretion. Mouse–cockroach interactions also changed following exposure of the mouse to the spray, as the mouse incited fewer interactions and more frequently struck with its forepaws than with its mouth. Monomorium sp. and Camponotus sp. ants also were repelled by the secretion, and Periplaneta americana were significantly irritated by the application of E. floridana secretion. The secretion is able to deter without physical contact, presumably via noxious volatiles. Eurycotis floridana is capable of accurately aiming the discharge as well as ejecting it several times the length of its body. It also was found that E. floridana exhibits autotoxic responses following contact with the secretion of conspecifics.     

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.     

Influence of water-stressed cassava on Phenacoccus herreni and three associated parasitoids

The influence of cassava Manihot esculenta Crantz grown under condition of water-stress on development and reproduction of the cassava mealybug, Phenacoccus herreni Cox & Williams, and levels of parasitism of three encyrtid parasitoids, Apoanagyrus diversicornis Howard, Aenasius vexans Kerrich, and Acerophagus coccois Smith, were studied in the laboratory. Two cassava cultivars were used: CM 507-37 (drought-tolerant) and CMC 40. A 30 day period of water stress, imposed by reducing the irrigation volume, led to a reduction in shoot development and stomatal conductance of leaves of both cassava genotypes. Phenacoccus herreni development and reproduction were favoured by cassava under water shortage. Parasitism decreased and water stress appeared to enhance the encapsulation of parasitoid eggs or larvae by the mealybug. In the case of the parasitoid A. diversicornis, there was a decrease in size of female progeny, suggesting a lower fitness in this species on cassava plants under water stress. All results indicated that cassava grown under low water availability favoured P. herreni development and reproduction, and affected the success of parasitism and, depending on the species, parasitoid development. The drought-tolerance characteristic of cassava genotypes and parasitoid species most suitable for controlling P. herreni in drought-stricken areas are discussed.