Cardinium is associated with reproductive incompatibility in the predatory mite Metaseiulus occidentalis (Acari: Phytoseiidae)

Cardinium, a bacterium from the Bacteroidetes group, is associated with reproductive manipulations such as cytoplasmic incompatibility, parthenogenesis, and feminization in some arthropod species. We, and others, have shown that Cardinium, but not Wolbachia, is an endosymbiont in some populations of Metaseiulus occidentalis, a phytoseiid mite that is an important predator of spider mite pests of agricultural crops. However, the precise biological effects that Cardinium may have on M. occidentalis remain unclear. In this study we show, in two sets of crosses between different colonies of Cardinium-free (C-) M. occidentalis females and Cardinium-containing (C+) males, that fecundity was reduced in parental females, F1 progeny survival rates were reduced, and fewer female progeny were produced when compared to the reciprocal and control crosses. There were no differences in these attributes in the reciprocal and two control crosses. Cardinium was transmitted maternally but there was no observed paternal transmission. Finally, Cardinium did not cause asexual (thelytoky) reproduction in M. occidentalis. Thus, Cardinium is associated with nonreciprocal reproductive incompatibility in M. occidentalis and our results support the hypothesis that Cardinium is a reproductive parasite in this agriculturally important predator. Cardinium may therefore affect the evolution and ecology of M. occidentalis and biological control efforts using this mite.     

Morphological and molecular characterization of a new microsporidian species from the predatory mite Metaseiulus occidentalis (Nesbitt) (Acari,Phytoseiidae)

A new microsporidian species is described from the predatory mite Metaseiulus (formerly Typhlodromus or Galendromus) occidentalis (Nesbitt) (Acari, Phytoseiidae). The ultrastructure of this new species is presented together with the first molecular characterization for a microsporidium of mites. All stages of this new microsporidium are haplokaryotic and develop in direct contact with the host-cell cytoplasm. Sporogony is disporoblastic and spores are formed in eggs, immature stages, and adults of M. occidentalis. There are two morphological classes of spores, one with a short polar filament (3-5 coils) that measured 2.53 x 1.68 microm and one with a longer polar filament (8-9 coils) that measured 3.14 x 1.77 microm. Horizontal transmission of this new species occurs by cannibalism of eggs and other stages and perhaps involves the spores with the long polar filament. Spores with the short polar filament may play a role in autoinfection and vertical (transovarial) transmission that is highly efficient in transferring the microsporidium from adults to progeny. Analysis of the small subunit ribosomal DNA indicated that this species from M. occidentalis is most closely related to the Nosema/Vairimorpha clade of microsporidia. A conflict between the morphological and molecular data is discussed. The species is compared to previously described microsporidia of arachnids resulting in creation of Oligosporidium occidentalis n. sp. in the family Unikaryonidae.     

The predatory mite Metaseiulus occidentalis: mitey small and mitey large genomes

Metaseiulus occidentalis is a representative of an important family of mites (Arthropoda: Chelicerata: Acari: Phytoseiidae) that are effective predators of pest mites in agricultural crops around the world. Like many arthropods, this mite contains multiple genomes, including the genomes of several microbial symbionts as well as its own mitochondrial and nuclear genomes. The mitochondrial genome is “mitey” large at 25 kb, due to duplication and triplication of genes. By contrast, the nuclear genome is “mitey” small at 88 Mb. This mite has a parahaploid genetic system, tolerates inbreeding, and has a haploid chromosome number of 3. This predator was genetically improved for use in agriculture by developing strains that lacked the ability to overwinter in diapause or were resistant to multiple pesticides, and can be genetically modified using recombinant DNA methods. Sequencing the nuclear genome would provide useful insights that could enhance genetic improvement programs that would result in improved pest management, could provide genes needed to resolve the evolutionary relationships of this family, and could serve as a model for understanding the evolution and genetics of chelicerate arthropod predators.     

Diapause behavior of genetically-improved strains of the spider mite predator Metaseiulus occidentalis (Acarina: Phytoseiidae)

Diapause behaviou of two strains of Metaseiulus occidentalis (Nesbitt) was compared as part of a project to evaluate this predator as a control agent for Tetranychus urticae Koch on roses in greenhouses in Berkeley, California. One strain had a normal diapause and the other had been genetically selected for non-diapause. Predators in diapause sought shelters on a simple bean plant test system. Females lacking the ability to diapause and the normal strain under long photophases tended to disperse aerially from the bean plant systems, particularly when prey were scarce. M. occidentalis females with the capacity to diapause reared near the critical photophase (11.2h) at fluctuating temperatures of 17.5 to 24.5°C within a 24 h period were influenced by the availability of prey: if prey were lacking, predators were more likely to enter diapause than if prey were abundant.Starved females left the plant system, probably through aerial dispersal, and mated females dispersed more readily than males and virgin females under long daylengths. No differences in aerial dispersal tendency were observed at short photophases. Aerial dispersal of adult females occurred at air speeds of ca. 1.5 m/s.

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Comportement lors de la diapause de souches génétiquement sélectionnées du prédateur Metaseiulus occidentalis (Acari; Phytoseiidae)

Résumé Au cours d'un programme destiné à évaluer la valeur du prédateur, Metaseiulus occidentalis Nesbitt pour lutter contre Tetranychus urticae Koch sur roses dans les serres de Berkeley (California) nous avons comparé chez deux souches le comportement lié à la diapause. Une souche a une diapause normale, l'autre a été sélectionnée génétiquement pour son absence de diapause. Les adultes avec diapause normale recherchent des abris sur un dispositif expérimental conçu à partir d'un pied de haricot. Les femelles perdant l'aptitude à la diapause ainsique celles de la souche normale soumise à une longue photophase ont tendance à se disperser par voie aérienne à partir du dispositif expérimental, surtout quand les proies sont rares.Les femelles de M. occidentalis aptes à la diapause élevées aux environs de la photophase critique (11.2h) avec des températures variant entre 17°5 et 24°5 pendant le nyctémère réagissent en fonction de la disponibilité en proies: quand les proies manquent, les prédateurs ont plus tendance à entrer en diapause que quand les proies sont abondantes.Avec une longue photopériode, les femelles à jeûn abandonnent le système expérimental, (probablement par dispersion aérienne) et les femelles fécondées se dispersent plus facilement que les mâles et les femelles vierges. Aucune différence dans la tendance à la dispersion aérienne n'a été observée aux courtes photophases. La dispersion aérienne des femelles adultes s'est produite à des vitesses de 1.5 m/sec.

     

Heat curing Metaseiulus occidentalis (Nesbitt) (Acari,Phytoseiidae) of a fitness-reducing microsporidium

Laboratory colonies of the predatory mite Metaseiulus occidentalis in Gainesville, FL were found to be infected with an undescribed microsporidium. Experiments were performed to quantify the effect of infection on the fitness of M. occidentalis and to determine if heat treatment can cure mites of the microsporidium. The colonies tested were derived from an isofemale line so that differences in performance could be attributed to the presence of microsporidia. A subcolony of an uninfected isofemale line was infected with the microsporidium by feeding females infected eggs from another colony of M. occidentalis. Infected mites had a shorter mean (+/-SD) female life span (7.4 +/- 2.9 vs. 10.0 +/- 2.8 days), lower mean oviposition (1.6 +/- 0.7 vs. 2.2 +/- 0.4 eggs/day), and a male-biased sex ratio (43 +/- 16% vs. 57 +/- 15% female progeny). The infection was reduced temporarily in colonies initiated from mites that were reared in a growth chamber at 33 degrees C from egg to adult, but healthy colonies only were established from the progeny of the heat-treated adults. These colonies remained free of infection for 10 weeks.     

Heterochromatinization,chromatin elimination and haploidization in the parahaploid mite Metaseiulus occidentalis (Nesbitt) (Acarina: Phytoseiidae)

Embryogenic mitoses, mitoses in females and spermatogenesis are described in the predatory mite Metaseiulus occidentalis (Nesbitt). At 22° C, egg development lasts approximately 4 days. Six chromosomes are seen in mitotic metaphases and anaphases of 0–24 h eggs. Toward the end of this period some embryo squashes have patches of cells containing nuclei which are partially heteropycnotic. These patches of cells apparently increase in size with the age of the embryo. In approximately 1/2 of all 24–48 h-old eggs they encompass all or most cells of the embryo. In these embryos metaphases involved 6 chromosomes, anaphases 3. Either prior to, or following metaphase, a pairing of chromosomes appeared to take place to form 3 units which resembled meiotic diplotene chromosomes where there is opening out between homologues. At metaphase, two sets of 3 chromosomes were slightly differentially stained. One, designated the H set, was darker and slightly more contracted than the other, the E set. At anaphase, 3H and 3E chromosomes segregated in a reductional division retaining the differential contraction until telophase. No cytokinesis appeared. The H set appeared to remain contracted while the E set decontracted to assume the appearance of an interphase nucleus. Both of these entities, side-by-side, created the partially heteropycnotic nucleus mentioned above. The H set then appeared to be excluded from the cell. Mitotic meta and anaphases involving 6 chromosomes were noted in female deutonymphs. Spermatogenesis appeared to encompass an equational division of 3 chromosomes, with the formation of a binucleate spermatid. Two tail structures appeared juxtaposed at the edge of each spermatid and thereafter a separation into two individual sperms occurred. —While mitosis was not studied in known males, we believe that the embryos exhibiting heterochromatinization and elimination of chromosomes in most or all cells were in fact demonstrating parahaploidization.This paper is dedicated to the memory of Professor S.W. Brown and is presented to Professor H. Bauer in honor of his 75th birthday     

Differential impact of egg predation by Zetzellia mali (Acari: Stigmaeidae) on Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

The differential impact of Zetzellia mali on the phytoseiids Metaseiulus occidentalis and Typhlodromus pyri was studied in the laboratory and by analysis of population from orchard plots that contained either phytoseiid, similar numbers of prey mites and high or low densities of Z. mali. Five hypotheses were evaluated to explain why Z. mali had more impact on M. occidentalis in the field than on T. pyri. Given equal opportunity, Z. mali adult females did not consume more M. occidentalis eggs than T. pyri eggs nor did adult females of either phytoseiid inflict greater mortality on Z. mali eggs or larvae through attack or consumption. There was no difference in the within-tree association of Z. mali adult females with eggs of either phytoseiid species nor were there differences in the way prey mites (all life stages) were spatially partitioned between adult female Z. mali as compared with adults and deutonymphs (combined) of either phytoseiid. The foraging area of adult female Z. mali and the oviposition locations of the two phytoseiids from both field and laboratory data were compared using spatial statistics. Metaseiulus occidentalis laid significantly more eggs in the primary foraging area of adult female Z. mali than T. pyri. This was the only factor identified which may explain the greater impact of Z. mali on M. occidentalis. The impact of these interspecific effects on the persistence of predatory mite guilds and biological control are discussed.     

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

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

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

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

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

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

Molecular and morphological boundaries of the predatory mite Neoseiulus californicus (McGregor) (Acari: Phytoseiidae)

Neoseiulus californicus (McGregor) is a natural enemy of pest mites used worldwide in many crops. Its correct identification is thus essential to ensure biological control success. The present study aimed to characterize molecular and morphological intraspecific variations for assisting in the diagnosis of the species and to build baseline information about expected variations within a commercially important phytoseiid species. Morphological and molecular [12S rRNA, cytochrome b mitochondrial (mt)DNA, and internal transcribed spacer] analyses were carried out on fourteen populations collected worldwide and on one mass‐reared strain. The genetic distances between the specimens of N. californicus and another related species were high and no overlap was observed, sustaining the reliability of such molecular methods for assisting a specific diagnosis. Furthermore, the genetic distances between populations of N. californicus were very low and overlap between intra‐ and interpopulations distances was observed, except for two populations collected in France (Marsillargues and Midi‐Pyrénées). The high mitochondrial differentiation between these two latter populations and the others questions their specific status: do they belong to the species N. californicus or to another cryptic species? However, using nuclear DNA marker analyses, no distinct differences were observed. Furthermore, even if significant morphological differences were observed between the populations, these differences were very small and the standard errors within each population were very low. We thus concluded that all the populations studied belong to the species N. californicus, despite unexpected high mitochondrial variations. The present study thus displays the importance of an integrative taxonomic approach for avoiding misidentifications. A discussion on morphological and mtDNA variations in relation to diagnostic reliability is developped. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 393–406.     

Side-effects of three pesticides on the predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae)

Side-effects of three commonly used pesticides in Iran were evaluated on an introduced strain of the predatory mite. Phytoseiulus persimilis Athias-Henriot, reared for about 10 years without exposure to any pesticides. Application of pesticides was carried out either to detached bean leaves using a Potter tower at 1 mg wet deposit per cm2 or by a hand sprayer on bean plants until run off. According to an EPPO decision making scheme, pirimiphos-methyl was found to be harmful (E=90.8%) and heptenophos harmless (E=-3.7%) to the predatory mite in the residual initial toxicity tests. For determination of the hazard class of malathion a field test was found to be necessary (E=59.8%). Categories of 1, 2, 3 were determined for heptenophos, malathion and primiphos-methyl, respectively, using IOBC classification. Despite being harmful, it is possible to use pirimiphos-methyl 10 days before release of P. persimilis. Investigation of the contribution of both lethal and sub-lethal effects to total impact indicated the dominance of lethality in the case of pirimiphos-methyl, while malathion acted by both mechanisms. Heptenophos did not have negative effects on fecundity of P. persimilis but rather caused a higher rate of fecundity in comparison with the control. The mortality found in the heptenophos test was not significantly different from the control.     

Loss of sex-allocation plasticity in the predatory mite Metaseiulus occidentalis and possible triggering cues

In two recent studies, we showed that females from a long-established Metaseiulus (Typhlodromus or Galendromus) occidentalis laboratory colony reared under crowded conditions for approximately 27 years produced equal numbers of male and female progeny. This contrasts with our previous results that showed the females from the same colony produced progeny with a female-biased sex ratio. These discrepant findings raised the possibility that dense rearing conditions can affect the sex-allocation plasticity in this important biological control agent. In the current study, we show that females from a newly collected M. occidentalis colony initially were capable of producing progeny with a female-biased sex ratio, but lost this ability over a 12-month period (43 generations) when reared under dense rearing conditions and provided with damaged prey. We demonstrate that females from this newly collected colony initially could adjust offspring sex ratio in response to both conspecific cues and cues produced by their damaged prey. The results suggest that dense rearing conditions, which are often used in commercial rearing facilities, may have detrimental effects on the reproductive fitness of M. occidentalis and, possibly, other predatory mites.     

Effect of several different pollens on the bio-ecological parameters of the predatory mite Typhlodromus athenas Swirski and Ragusa (Acari: Phytoseiidae)

The development, survivorship, and reproduction of the predacious mite Typhlodromus athenas Swirski and Ragusa were studied in the laboratory by rearing the predator on nine different plant pollens [almond(Prunus amygdalis Batsch), apple (Malus domestica Borkh.), apricot (Prunus armeniaca L.), cherry (Prunus avium L.), pear (Pyrus communis L.), plum (Prunus domestica L.), walnut (Juglans regia L.), olive (Olea europaea L.), Typha sp.], and pollen collected from bee hives. All experiments were conducted in environmental chambers at 20 ± 1°C, 65% RH, and a photoperiod of 16:8 (L:D) h. Survival during immature development ranged from 81.1 to 96.0%. The shortest mean developmental time from egg to adult with respect to the range of pollen species was recorded for females and males fed on almond pollen (10.76 ± 0.18 and 10.45 ± 0.21 d, respectively), while the longest was on beehive pollen (26.97 ± 0.23 and 24.00 ± 0.25 d for females and males, respectively). Female longevity varied from 51.63 ± 5.52 d (olive pollen) to 102.81 ± 6.60 d (pear pollen), while fecundity ranged from 5.33 ± 2.35 eggs per female (beehive pollen) to 26.43 ± 1.73 eggs per female (almond pollen). The diet consisting of almond pollen resulted in the highest intrinsic rate of natural increase (r(m)) (1.00 d(-1)) and pollen collected from bee hives resulted in the lowest (0.013 d(-1)). These results showed that various pollen could favor the development of T. athenas, and also support the view that alternative food resources may play an important role in the field for sustaining and increasing the predator's population.     

Adaptability and efficacy of transgenic and wild-type Metaseiulus occidentalis (Acari: Phytoseiidae) compared as part of a risk assessment

Comparisons between transgenic (T) and wild-type Metaseiulus occidentalis colonies (COS) were made under laboratory conditions as part of a risk assessment effort prior to proposed field releases. There were no differences between the transgenic T18 colony and the COS strain in the daily egg production, hatchability at three temperatures and four relative humidity (RH) conditions, diapause incidence, or proportion of female progeny produced. Metaseiulus occidentalis eggs do not hatch at 38°C under any RH tested, nor at 33.5°C under 100% RH, indicating that high temperature and extreme RH affect egg hatch negatively. At 28.5 and 33.5°C, fewer eggs hatched at 32.5% than at 75.5 and 93% RH. Metaseiulus occidentalis cannot survive on any plant tested without prey nor on a diet of pollen alone; adult female mites cannot prey or survive on a diet of eggs and larvae of two lepidopteran species, indicating that the suitability of food sources has not been altered in the transgenic strain. Two subcolonies, derived from two transgenic strains using single females, differed in the rate of egg hatch at 28.5°C under 32.5 and 100% RH, indicating that reduced genetic variation and/or random genetic drift in the two lines may have led to differences in some biological characters. Since we did not find any significant differences between the T18 and COS colonies in the traits tested, the T18 colony is not expected to exhibit any new biological attributes in a proposed short-term field release.     

A novel disease affecting the predatory mite Phytoseiulus persimilis (Acari, Phytoseiidae): evidence for the involvement of bacteria

Adult female Phytoseiulus persimilis Athias-Henriot (Acari, Phytoseiidae) of a laboratory population show drastic changes in foraging behavior, anatomy and life history compared to typical laboratory populations. We demonstrated earlier that the set of characteristic symptoms, called non-responding (NR) syndrome, is transmitted horizontally between and among predator generations via feces and debris deposited by symptomatic females. Here, we prove that bacteria present in feces and debris deposited by symptomatic females are involved in the induction of the NR-syndrome. The potential of predator products to induce the NR-syndrome was assayed by keeping healthy adult female predators during a period of 3 days on prey-infested bean leaves, which had previously been sprayed with an aqueous suspension of feces and debris. The NR-syndrome was clearly induced in those predators that had been exposed to a suspension collected from symptomatic females (incubation time 4-6 days, 93% shrunken females), whereas predators exposed to a suspension collected from non-symptomatic females did not show the NR-syndrome. Moreover, predators from the first group transmitted infectious products themselves already 5 days after the initial exposure, whereas this was not the case for the second predator group. The bioassay used in the present study is important for laboratories and companies as it can be applied for testing the presence of the novel disease in populations of P. persimilis. To investigate the involvement of bacteria in syndrome induction we (1) eliminated bacteria from a feces-and-debris suspension of symptomatic females by passing the suspension through a bacterial microfilter and (2) added the antibiotic tetracycline to a suspension of feces and debris from symptomatic females. A suspension of feces and debris collected from symptomatic females did not induce the NR-symptom after bacteria had been eliminated, whereas an untreated portion of the same suspension did so. Moreover, the NR-syndrome was induced in predators exposed to an aqueous suspension of the residues that had not passed the bacterial filter. A suspension of feces and debris collected from symptomatic females, to which the antibiotic tetracycline had been added, did not induce the NR-syndrome whereas the same suspension did induce all symptoms when no tetracycline was added. These findings prove that bacteria are involved in the induction of the NR-syndrome. The results are discussed in the context of mite pathology and biological control.     

The effects of prestarvation diet on starvation tolerance of the predatory mite Neoseiulus californicus (Acari: Phytoseiidae)

Understanding the factors affecting stress tolerance in phytoseiid mites is critical for their integration into biological control programs. In the present study, the effects of diet (varying in prey species, physiological status and phenotype) are examined on the future starvation tolerance of the predatory mite Neoseiulus californicus McGregor (Acari: Phytoseiidae). The predators are fed from egg to adulthood on diapausing or nondiapausing Tetranychus urticae, Tetranychus kanzawai (wild and albino strains) or the nondiapausing species Panonychus citri (wild and albino strains). Thereafter, 3‐day‐old mated adult females are held without food at 25 ± 1 °C and a relative humidity of 98 ± 2%. The survival of these starved females is observed daily until all females have died. The survival curves and mean survival times of N. californicus are found to vary among prey types and are significantly longer when the predator is fed with diapausing prey. This enhanced survival is consistent with high concentrations of glycogen and triacylglyceride in the body of the predator at the onset of starvation. The predators fed nondiapausing prey have shorter survival times, and the glycogen and triacylglyceride contents in their bodies are low or undetectable. The protein contents of the predator's body are similar after consuming different prey types, except for a high concentration when fed the albino strain of P. citri. Protein content is unlikely to play a direct role in starvation tolerance, although it may affect the response to varying glycogen and triacylglyceride levels. These findings indicate that nutritional value of prey has a strong impact on the starvation tolerance of N. californicus.