Photoperiod and temperature studies to determine whether diapause is found in successive generations of the African phytoseiid,Euseius fustis (Pritchard and Baker) (Acari: Phytoseiidae)

Photoperiod and temperature conditions known to induce diapause in tropical arthropods were tested on two generations (G₀ and G₁) of the phytoseiid, Euseius fustis. Failure to lay eggs or a pre-oviposition period longer than 15 days were the criteria used to determine whether females were in diapause. Females reared from egg to adult and held throughout adult life under cyclic temperatures of 29/20°C in combination with long photophases of 16L: 8D and 14L: 10D showed no indication of aestival diapause. Similarly, hibernal diapause was not induced in females reared under a constant temperature of 18°C and a photophase of 8L: 16D. Under the various test conditions, females initiated oviposition within an average of 4 days. Overall, pre-oviposition patterns for G₀ and G₁ females were similar under the same test conditions. Reproductive patterns based on the mean number of eggs per female per day varied only slightly between generations for the same treatments. No behavioural or morphological attributes associated with diapause were observed.     

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.     

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.     

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.     

Development, survival and reproduction of Euseius finlandicus (Acari: Phytoseiidae) at different constant temperatures

Development, survival and reproduction of Euseius finlandicus Oudemans were studied at seven constant temperatures (15, 20, 25, 27, 30, 32 and 34°C) in the laboratory. Within the temperature range tested, developmental period from egg to adult varied from 148 to 360.5h and 133.7 to 336.5h for females and males, respectively. The lower thermal threshold for immature development for females and males was 8.9 and 6.4°C, respectively. Survival during immature development exceeded 90% at all the temperatures from 15 to 32°C, but at 34°C an abrupt decline was recorded. Female longevity decreased gradually from 82.7d at 15°C to 12.2 d at 34°C. The mean generation time ranged from 44.3d at 15°C to 15.9d at 32°C. The highest r _m value (0.2817) was obtained at 30°C and the lowest at 15°C (0.0976). Temperatures above 30°C had an adverse effect on population increase.     

Prey stage preference and functional response of Euseius hibisci to Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae)

The aims of this study were: (a) determine the prey stage preference of female Euseius hibisci (Chant) (Phytoseiidae) at constant densities of different stages of Tetranychus urticae Koch (Tetranychidae), (b) assess the functional response of the predator females to the varying densities of eggs, larvae, or protonymphs of T. urticae, and (c) estimate the functional response of E. hibisci when pollen of Ligustrum ovalifolium was present as well. We conducted experiments on excised pieces of strawberry leaf arenas (Fragaria ananassa) under laboratory conditions of 25 ± 2 °C, 60 ± 5% RH and 12 h photophase. Our results indicated that the predator consumed significantly more prey eggs than other prey stages. Consumption of prey deutonymphs and adults was so low that they were excluded from the non-choice functional response experiments. The functional response on all food items was of type II. The two parameters of the functional response were estimated for each prey type by means of the adjusted non-linear regression model. The highest estimated value a (instantaneous rate of discovery) and the lowest value of T_h (handling time, including digestion) were found for the predator feeding on prey eggs, and a was lowest and T_h highest when fed protonymphs. Using the jack-knife method, the values for the functional response parameters were estimated. The values of a and T_h produced by the model were similar among all prey types except for the eggs, which were different. Using pollen simultaneously with prey larvae decreased the consumption of the latter over the full range of prey densities The suitability of this predator for biological control of T. urticae on strawberry is discussed.     

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.     

Pearl bodies of Cayratia japonica (Thunb.) Gagnep. (Vitaceae) as alternative food for a predatory mite Euseius sojaensis (Ehara) (Acari: Phytoseiidae)

On the young leaves, shoots, and buds of Cayratia japonica (Thunb.) Gagnep. (Vitaceae), we observed nutritious bodies called pearl bodies and hypothesized that they are utilized by generalist predators as alternative foods. Some ambulate organisms consume pearl bodies in the wild and the predatory mite Euseius sojaensis (Ehara) (Acari: Phytoseiidae) was considered as a primary candidate. Pearl bodies promoted E. sojaensis settlement on C. japonica leaves and E. sojaensis could prey on the phytophagous mite Tetranychus kanzawai Kishida (Acari: Tetranychidae) when the predators settle on a leaf before the prey. In addition, the presence of pearl bodies did not reduce predation of E. sojaensis on T. kanzawai. This was seemingly because food quality of T. kanzawai was higher than pearl bodies. These results implied that pearl bodies on C. japonica leaves are utilized by E. sojaensis as alternative foods.     

Survival,development and reproduction of Euseius finlandicus (Oudemans) (Acari: Phytoseiidae) fed on various kinds of food substances

Laboratory experiments were conducted to investigate the effect of diet on the biology of the phytoseiid mite, Euseius finlandicus (Oudemans). The predatory mite was able to develop and reproduce better when fed on the eriophyid mites, Aceria olivi (Zaher and Abou-Awad), Aceria dioscoridis (Soliman and Abou-Awad) and Cisaberoptus kenyae (Keifer). The developmental time of immature stages was the shortest when fed on motile stages of eriophyid mite species, followed by the spider mite, Tetranychus urticae Koch, and then pollen grains of Ricinus communis L., Phoenix dactylifera L. and Helianthus annuus L. Total egg production was highest when the predator fed on A. olivi, A. dioscoridis and C. kenyae recording at the rate of 51.0 50.0 and 43.84 eggs/female, respectively, but lowest on pollen grains, R. communis, P. dactylifera and H. annuus at the rate of 11.96, 5.3 and 2.0 eggs/female, respectively. But, the reproduction was nil on the tetranychid mite, T. urticae. Also, sex ratio of the progeny favoured females, when the predatory mite was reared on the eriophyid preys. E. finlandicus recorded the highest intrinsic rate of increase (r_m?=?0.31 females/female/day) when fed on A. dioscoridis, followed by (0.30 and 0.23 females/female/day) when fed on A. olivi and C. kenyae, respectively. In contrast, the lowest intrinsic rate of increase (r_m?=??0.31) was noted when fed on H. annuus pollen grains. The eriophyid mite, as a prey, recorded the shortest developmental time and highest oviposition rate of E. finlandicus.     

Overwintering sites and winter mortality of Euseius finlandicus (Acari: Phytoseiidae) in a peach orchard in northern Greece

In peach orchards of northern Greece Euseius finlandicus Oudemans overwinters in various sites on the trees, usually in groups of 5–15 females. Overwintering females were mostly found in bark crevices near the hibernation cocoons of the peach moth Adoxophyes orana (Fischer von Rosslerstamm), and also in small crevices of the bark and in pedicels left after fruit harvesting. Empty scales of dead diaspidids and dead coccids, cocoons of lacewings and mummies of parasitized aphids sporadically found on trees, were less common overwintering sites for females of the mite. In two successive years, mortality of overwintering females was very low, approximately 4 and 4.5% which could be due to the mild climate of northern Greece and the high cold tolerance of the mite. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Effect of relative humidity and origin of isolates of Neozygites tanajoae (Zygomycetes: Entomophthorales) on production of conidia from cassava green mite, Mononychellus tanajoa (Acari: Tetranychidae), cadavers

Neozygites tanajoae is a very specialized fungus pathogenic to the cassava green mite (CGM), Mononychellus tanajoa, an important cassava pest introduced to Africa from the Neotropics. Conidial discharge from cadavers of CGM that died from infections with 14 isolates of N. tanajoae collected from diverse climates of Brazil was quantified to help select potential candidate strains for introduction to Africa. Studies aimed to identify isolates with lower requirements for relative humidity for sporulation and isolates that discharged more conidia during short periods of moisture. At 96 ± 0.5% RH, production of conidia was variable and even isolates from the Brazilian semi-arid region, e.g., Petrolina and Itaberaba, produced few conidia. Significant differences in the numbers of conidia produced by diverse Brazilian isolates were observed after 6, 9 and 12 h at 100% RH. At 100% RH, production of primary conidia increased considerably from an average of 57 ± 4 conidia at 6 h to 509 ± 37 conidia at 12 h. The isolate sporulating least (BIN21) discharged only 45.7% of the number of conidia produced by isolate BIN1, one of the isolates producing the most spores. Results from this study demonstrate that differences in production of conidia among isolates should be considered when selecting Neozygites isolates for new biological control introductions.     

有益真绥螨与巴氏新小绥螨的集团内捕食和同类相残作用

【目的】有益真绥螨Euseius utilis是北方地区广泛分布的一种多食性植绥螨,而巴氏新小绥螨Neoseiulus barkeri目前在我国广泛应用于农业害虫的生物防治中。本研究旨在对巴氏新小绥螨在本地的应用进行风险评估及为与有益真绥螨的联合释放提供依据。【方法】本研究在室内通过一系列实验,比较了实验室饲养的巴氏新小绥螨和采自内蒙古农业大学校园的有益真绥螨的雌成螨对同种或异种未成熟螨的捕食量、存活时间及产卵量的差异,检测了有益真绥螨与巴氏新小绥螨两种植绥螨相互之间的攻击强度以及种内和种间的相互作用。【结果】两种植绥螨都难以刺吸同种或异种植绥螨的卵,而对同种或异种植绥螨幼螨的捕食量最大,其次是对若螨。在无共同食物的情况下,有益真绥螨雌成螨对同种植绥螨幼螨的攻击性比对异种植绥螨幼螨的攻击性强(BreslowDay检验:χ~2=13.84,df=1,P0.001),且有益真绥螨对同种植绥螨幼螨的捕食量(9.10±1.65头)高于对异种植绥螨幼螨的捕食量(5.31±1.43头)(T检验:t=5.487,P0.001),巴氏新小绥螨对异种植绥螨幼螨的捕食量(7.48±0.75头)高于对同种植绥螨幼螨的捕食量(4.75±0.58头)(T检验:t=9.110,P0.05)。【结论】有益真绥螨更倾向于捕食同种幼螨而发生同类相残;巴氏新小绥螨更倾向于捕食异种幼螨而发生集团内捕食。     

Predatory behaviors of Neoseiulus californicus and Galendromus helveolus (Acari: Phytoseiidae) attacking Oligonychus perseae (Acari: Tetranychidae)

Predatory behaviors of Neoseiulus californicus (McGregor) and Galendromus helveolus (Chant) attacking Oligonychus perseae Tuttle, Baker and Abbatiello on avocado leaves were videotaped and analyzed. Behaviors were recorded for “fresh” predators that were used ≤ 48 hr post receipt from a commercial insectary and “cold stored” predators that were maintained at 12°C for ≈ 14 days. Fresh and cold stored G. helveolus were observed to attack O. perseae only after invading webbed nests. Conversely, fresh and cold stored N. californicus employed three different modes of predatory attack: (1) intercepting and attacking migrant O. perseae outside of web nests; (2) attacking prey through nest webbing; or (3) invading and attacking O. perseae inside nests. Predatory efficacy of both N. californicus and G. helveolus was reduced following cold storage, as both species engaged in certain predatory behaviors less frequently in comparison to predators that were not stored at low temperatures. Our observed results for N. californicus and G. helveolus attacking O. perseae are interpreted in relation to the chaetotaxy hypothesis, which proposes that phytoseiid invasion efficiency and propensity of webbed nests is facilitated by dorsal setal lengths. This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparison of biological and demographic parameters forMononychellus progresivus andOligonychus gossypii on cassava: influence of temperature

The effect of five constant temperatures (16, 22, 26, 31 and 36°C) on biological (survival and duration of developmental stages, fecundity and longevity of females, sex-ratio) and demographic parameters (R _o , G, r _m and λ) of the two main phytophagous mites that attack cassava in Africa,Mononychellus progresivus Doreste andOligonychus gossypii (Zacher), was studied in the laboratory. Experiments were performed simultaneously on the two mite species reared on the same cassava variety (1M20) under controlled conditions: 70±10% r.h. and 12L∶12D. The lower thermal threshold was 13°C forM. progresivus and 11°C forO. gossypii. Both species developed in the range from 22 to 36°C. The shortest development time was obtained at 31°C for both species; it was 7.2 days forM. progresivus and 8.2 days forO. gossypii. Maximum fecundity of both species was recorded at 26°C with 42.1 eggs forM. progresivus and 36.3 eggs forO. gossypii The highest intrinsic rate of increase (r _m ) was obtained at 31°C for both species with 0.289 and 0.214 forM. progresivus andO. gossypii, respectively.     

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.     

Effect of mating frequency on fecundity and longevity of the predatory mite <Emphasis Type="Italic">Kampimodromus aberrans</Emphasis> (Acari: Phytoseiidae)

The effect of single versus multiple mating on longevity and fecundity as well as the number of matings required to maximize a female’s reproductive success of the predatory mite Kampimodromus aberrans Oudemans were studied under laboratory conditions. Newly emerged adult females of the stock colony of K. aberrans were placed individually on a bean leaf disc, and maintained at 25°C and 16:8 LD. A young male remained with a female for limited periods or continuously. Mating was a requisite for oocyte maturation and oviposition. Females which mated three to four times during their life and females in continuous presence of males, laid significantly and considerably more eggs than single-mated females. Virgin females lived the longest, and those in continuous presence of males the shortest. In all cases and irrespective of the number of matings, the sex ratio of the offspring was male-biased in the first three to four days of oviposition period, and female-biased in later days.     

Influence of selected fruit tree pollen on life history of <Emphasis Type="Italic">Euseius stipulatus</Emphasis> (Acari: Phytoseiidae)

Euseius stipulatus (Athias-Henriot) is a predatory mite widespread in the Mediterranean region considered to be important for the biological control of spider mites in citrus orchards. Development, survival and reproduction of this phytoseiid mite feeding on seven commercially obtained pollen were studied under constant laboratory conditions (20 ± 1°C, RH 65 ± 5%, photoperiod 16L: 8D h). Mites were kept individually at rearing units with ample quantity of almond (Prunus amygdalus Batch), apple (Malus domestica Borkh), apricot (Prunus armeniaca L.), cherry (Prunus avium L.), pear (Pyrus communis L.), plum (Prunus domestica L.) and walnut (Juglans regia L.) pollen as food source. Developmental time from egg to adult varied between the several pollen tested from 8.38 ± 0.08 to 9.58 ± 0.11 days for females and from 8.23 ± 0.12 and 9.07 ± 0.12 days for males. Female longevity varied from 11.53 ± 1.22 to 51.38 ± 2.45 days, while fecundity ranged from 22.84 ± 2.30 to 43.61 ± 3.78 eggs/female. The predator was unable to reproduce when feeding on walnut pollen. Data were submitted to life table analysis and values of the intrinsic rate of increase were derived, ranging from 0.079 to 0.146 (day⁻¹). The cumulative Weibull function that was used to describe the age specific survival of females produced excellent fits to the survival data. Results show that almond, plum, cherry and apricot pollen possess higher nutritional value for E. stipulatus than pear and apple pollen and thus may contribute in sustaining and increasing the predator population in field conditions. Walnut pollen can be utilized by the predator only to survive during short periods of time when principal or alternative food sources are scarce.     

Non-prey food for subsistence of Amblyseius idaeus (Acari: Phytoseiidae) on cassava in Africa

The nutritional effects of pollen from cassava, oil palm, castor bean, Leucaena leucocephala and a phloem exudate from cassava were tested in the laboratory for their effect on survivorship and oviposition of Amblyseius idaeus (Denmark & Muma). Survival and longevity of even-aged females were enhanced on the aforementioned non-prey diets when compared with the control of no food. Castor bean pollen provided sufficient nutrition to induce an irregular and low ovipositional rate of A. idaeus over a five week interval. Oviposition ceased after 48 h in the presence of the other non-prey foods or free-water only. These studies suggest that A. idaeus is capable of switching or supplementing its diet with non-prey foods during periods when spider mite populations are low.