Effect of prey density on sex ratio of two predacious mites, Phytoseiulus persimilis and Amblyseius womersleyi (Acari: Phytoseiidae)

The sex ratios of two phytoseiid mites, Phytoseiulus persimilis and Amblyseius womersleyi, were observed under various prey conditions. Upon consumption of abundant prey, both phytoseiids produced progeny in a female-biased sex ratio (approximately 0.8 females). When few prey were consumed, the sex ratio was lowered to 0.5 (the unbiased sex ratio). Under the conditions in which the unbiased sex ratio was observed, male and female progeny appeared in an alternating sequence. To determine the change in the sex ratio and the sequence of progeny, the size of eggs deposited by females under various prey conditions was first compared. Survivorship and developmental rate of progeny (immatures) hatched under ample and poor prey conditions were also examined. The eggs deposited under poor prey conditions were smaller than those deposited under ample prey conditions. This is an indication that the phytoseiid females did not invest extra energy into the eggs to secure survival of their progeny under poor prey conditions. The male and female progeny from the small eggs developed slowly, probably due to the small egg size. However, hatchability of the small eggs and survival of the immatures were the same as those of the normal eggs when the immatures were reared under ample prey conditions. The immature survivorship was little affected by the prey consumption rate of their mothers when the immatures were reared under poor prey conditions. We concluded that the sex ratio of phytoseiid mites is not determined by the characteristics of the progeny, but by the nutritious condition of the females.Exp Appl Acarol 22: 709723 © 1998 Kluwer Academic Publishers     

Effects of parent age at mating on reproductive response of Glyptapanteles flavicoxis (Hymenoptera: Braconidae), a larval parasitoid of the gypsy moth (Lepidoptera: Lymantriidae)

Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae) is a gregarious larval parasitoid of the Indian gypsy moth Lymantria obfuscata (Walker) (Lepidoptera: Lymantriidae), that is believed to have potential for inundative releases against gypsy moth populations, because it can be reared in large numbers with few hosts. Unfortunately, sex ratios in laboratory reared G. flavicoxis are usually male-biased, hindering efforts to mass release this species for biological control by making the production of females costly. Because parental age at time of mating is known to affect the sex ratio in some Braconidae, we crossed haploid males and virgin females at 0, 1, 4, 9, and 16 d old with at least 10 trials for each of the 25 combinations. Numbers and sex ratios of progeny produced by females each day were recorded. Both progeny and sex ratios (percentage of females) among progeny produced by ovipositing females of G. flavicoxis decreased markedly over time, so only the first days production need be used in mass rearing. The reduction in the proportion and numbers of females among progeny as females aged is consistent with sperm depletion. Approximately 30% of females in all age classes mated to newly emerged males (day 0) produced all male progeny, whereas only 10-15% of those mated to older males failed to produce any daughters. When crosses with only male progeny were excluded from the analysis, females mated to males 1 d old had higher sex ratios in progeny than those mated to males in other age classes. In addition, females mated the day that they emerged tended to have progeny with the highest sex ratios.     

Evaluation of the predacious mite Hemicheyletia wellsina (Acari: Cheyletidae) as a predator of arthropod pests of orchids

The cheyletid predator Hemicheyletia wellsina was found in association with orchid pests in a Florida greenhouse. The life history of H. wellsina was determined using the two-spotted spider mite, Tetranychus urticae, as prey in the laboratory. Hemicheyletia wellsina is arrhenotokous, with mated females producing a female-biased sex ratio of 3.4:1. Adult longevity averaged 30.2 days, with females producing an average of 18.5 eggs. Adult females live up to 17 days without food, surviving an average of 9.7 days. Hemicheyletia wellsina will readily feed on the phytoseiid Metaseiulus occidentalis, suggesting intraguild predation by H. wellsina on phytoseiids could be important in pest management programs. The average prey consumption of H. wellsina is 1.5 T. urticae females per day. The results of this study provide information on an under-studied group of mite predators. Although H. wellsina does not have the qualities that would be necessary for its use in an augmentative or classical biological control program, its use in naturally occurring biological control could be beneficial.     

Sex ratio variation in female-biased populations of Notostracans

Females from female-biased populations of the notostracan Triops newberryi produce viable eggs when reared in isolation. Clutches produced under such conditions exhibit sex-ratio polymorphism. One category of females (monogenics) produces only female offspring; the second category (amphigenics) produces clutches with a sex ratio of 3 females: 1 male. The relative proportions of the two categories of females varies significantly between populations and is correlated with population sex ratio. This correlation, and the pattern of offspring distribution in amphigenics, suggests that sex is determined by an autosomal Mendelian gene locus for which the male-determining allele is recessive. Limited pedigree analysis of lineages under selfing support the genetic model.     

Do components of colonization - dispersal cycles affect the offspring sex ratios of Banks grass mites (Oligonychus pratensis)?

The effects of two host plant species, immature rearing density, and feeding-induced leaf damage on the offspring sex ratios of individual Banks grass mites (BGM) (Oligonychus pratensis Banks) were examined. Offspring sex ratios did not differ among corn-reared females transferred to corn or two sorghum genotypes. Likewise, the offspring sex ratios did not differ among females from three rearing densities. However, female BGM reared on damaged leaves produced significantly more daughters than females reared on undamaged leaves. This suggests that offspring sex ratios of BGM change in response to a deteriorating food source. Greater daughter production on a deteriorating host results in more potential dispersers and colonizers of new host plants. This may be an important adaptation facilitating the colonization-dispersal episodes that characterize the BGM life history.     

Genetic and Environmental Factors Associated with Laboratory Rearing Affect Survival and Assortative Mating but Not Overall Mating Success in Anopheles gambiae Sensu Stricto

Anopheles gambiae sensu stricto, the main vector of malaria in Africa, is characterized by its vast geographical range and complex population structure. Assortative mating amongst the reproductively isolated cryptic forms that co-occur in many areas poses unique challenges for programs aiming to decrease malaria incidence via the release of sterile or genetically-modified mosquitoes. Importantly, whether laboratory-rearing affects the ability of An. gambiae individuals of a given cryptic taxa to successfully mate with individuals of their own form in field conditions is still unknown and yet crucial for mosquito-releases. Here, the independent effects of genetic and environmental factors associated with laboratory rearing on male and female survival, mating success and assortative mating were evaluated in the Mopti form of An. gambiae over 2010 and 2011. In semi-field enclosures experiments and despite strong variation between years, the overall survival and mating success of male and female progeny from a laboratory strain was not found to be significantly lower than those of the progeny of field females from the same population. Adult progeny from field-caught females reared at the larval stage in the laboratory and from laboratory females reared outdoors exhibited a significant decrease in survival but not in mating success. Importantly, laboratory individuals reared as larvae indoors were unable to mate assortatively as adults, whilst field progeny reared either outdoors or in the laboratory, as well as laboratory progeny reared outdoors all mated significantly assortatively. These results highlight the importance of genetic and environment interactions for the development of An. gambiae''s full mating behavioral repertoire and the challenges this creates for mosquito rearing and release-based control strategies.     

Sex ratios in natural populations ofAphelinus mali (Hym.: Aphelinidae) in relation to host size and host density

In an apple orchard at Armidale, the Northern Tablelands of NSW, population sex ratios ofAphelinus mali (Haldeman), an endoparasitoid of the woolly apple aphid,Eriosoma lanigerum (Hausmann) varied from 0.51 (proportion of males) at low host densities to female-biased at high host densities (proportion of males ranged from 0.35–0.39). This shift in sex ratio seems to be caused by the differences in allocation of sons and daughters to hosts of different sizes. In the fieldA. mali parasitizes all life stages (four nymphal instars and adult) of the woolly aphid upon encountering. According to Hughes'(1979) optimal diet model, such general host acceptance seems to be the best strategy. However, it allows the host nymphs or adults to continue to develop or reproduce until about to mummify (pupate). No mortality was observed when first or second-instar hosts were parasitized in the laboratory. Field collected small mummified hosts yielded male-biased sex ratios whereas large mummified hosts produced mainly females. In the laboratory, progeny from smaller hosts (first to third-instars) produced sex ratios which were not significantly different from 0.5 whereas progeny from larger hosts (third and fourth-instars) produced female-biased sex ratio. During winter (June–August) and early spring (September–October) when the host populations in the orchard were predominantly nymphs, the parasitoid tended to allocate equal resources to male and female offspring. In contrast, at peak population densities in summer and autumn (December–May) when larger hosts were available, the sex ratios were female-biased. The host size ofE. lanigerum andA. mali is, therefore, an important component in the dynamics of host-parasitoid interactions.     

Effect of mating combinations on the host parasitisation and sex allocation in solitary endoparasitoid,Aenasius arizonensis (Hymenoptera: Encyrtidae)

Aenasius arizonensis (Girault) is an important solitary endoparasitoid of Phenacoccus solenopsis Tinsley. To optimise the mass production of high-quality females, it is important to assess the influence of mating regimes on the progeny fitness and sex allocation. We, therefore, hypothesise that mating combinations in A. arizonensis adults emerged from different host instars may influence parasitism and sex allocation in the subsequent generation. Therefore, we compared three nymphal instars (1st, 2nd and 3rd) and adults host stages of P. solenopsis for parasitism and sex allocation by A. arizonensis. Further, F₁ female progeny of the parasitoid emerged from different host instars was henceforth evaluated for its fitness in six mating combinations. A. arizonensis females parasitised all the host stages except the 1st instar nymphs. The parasitised 2nd instar nymphs yielded only males, while the sex ratio in the later host instars was strongly female-biased. The parasitoid females preferred 3rd instar nymphs with respect to higher parasitism (74.0–84.0%) and produced more females in the F₁ progeny as compared to other host stages. F₁ females that emerged from 3rd instar nymphs produced significantly higher parasitism (74.0–79.0%). These mating combinations also yielded more female progeny in the F₂ generation. However, parasitism by F₁ females was significantly lower (9.0–12.0%) when mated with males that emerged from 2nd instar P. solenopsis nymphs. Moreover, latter combinations yielded only male progeny in F₂ generation. These findings can be used in laboratory mass rearing of this parasitoid vis-à-vis biological control of P. solenopsis.     

Diapause in the mite Metaseiulus occidentalis: Stages sensitive to photoperiodic induction

Stages of Metaseiulus occidentalis sensitive to photoperiod induction of diapause were determined by transferring various stadia into diapause-inducing conditions, and rearing them until adult females could be scored for reproductive condition. When eggs were transferred to 10 hr light at 19°C from 24 hr light at 25°C and the mites reared to adults, 92 per cent entered diapause. When larvae and all subsequent stages were kept under the inductive conditions, 62 per cent of adult females diapaused. Mites transferred as protonymphs into inductive conditions yielded only 10 per cent in diapause, and mites transferred as deutonymphs or newly emerged females did not enter diapause.However, adult females reared from eggs at 19°C under 12 hr light (which is near the critical photophase of 11·2 hr at 19°C) showed an unexpected sensitivity to photoperiod. Some newly emerged females oviposited upon transfer to an 8 hr photophase at 19°C. Some then stopped ovipositing and apparently entered diapause; these females resumed ovipositing after intervals ranging from 34 to 100 days. This was termed ‘switching’ into diapause. Some females reared under a 16 hr photophase at 19°C ‘switched’ also upon transfer as adults to shorter photophases—either 8 or 12 hr at 19°C. Thus, ‘switching’ may be due to transfer to shorter photophases. Promptness of mating vs delayed mating allowed ‘switching’ to be more easily detected.     

Experimental induction and termination of non-reciprocal reproductive incompatibilities in a parahaploid mite

Two experiments were conducted to investigate the relationship between rearing temperatures, the presence or absence of Wolbachia endosymbionts, and non-reciprocal reproductive incompatibility in inbred lines of the parahaploid phytoseiid mite Metaseiulus occidentalis (Nesbitt) (Acari: Phytoseiidae).Heat-treated females crossed with infected males reared at room temperature produced few eggs and high proportions of shriveled eggs. No female progeny were produced. The reciprocal cross was normal. A second experiment showed that the incompatible cross from the first experiment could be made compatible if the infected line was heat-treated and those males crossed with the original heat-treated females. Furthermore, a new incompatibility was induced in a formerly compatible cross when the newly heat-treated females were crossed with males from their base colony. Heat-treatment was correlated with the loss of Wolbachia in both experiments. Wolbachia may thus affect non-reciprocal reproductive incompatibility in M. occidentalis, and may produce a unique incompatibility phenotype in this parahaploid species, including both reduced numbers of male progeny and a lack of female progeny.     

Biological aspects and life table parameters of the predacious mite,Neoseiulus californicus (McGregor) (Acari: Phytoseiidae) consuming food types during immature stages and after adult emergence

To investigate the effect of food types on biological aspects and life table parameters of the predacious mite, Neoseiulus californicus (McGregor), immature stages were fed on pollen of maize, Zea mays L. and eggs of Tetranychus urticae Koch, while adult females were offered the same foods and the alternate shift of each food. All individuals of predator females reached adulthood when reared on maize pollen and prey eggs, but their life cycle was significantly longer on pollen than on prey. The shortest preoviposition period of N. californicus occurred after continuous feeding on T. urticae eggs, whereas the longest happened when fed on prey eggs switched to maize pollen. Continuous consuming of maize pollen showed the longest ovipositoin period for the predator, while rearing on maize pollen shifted to prey eggs exhibited the shortest duration. The longest longevity and highest sex ratio of females to total in N. californicus were recorded when continuously utilised pollen, whereas the shortest longevity and lowest sex ratio were noted when continuously consumed prey. On the contrary, the egg production and life table values of the predator were the highest by continuous feeding on prey eggs, while they were the lowest by continuous rearing on maize pollen. Therefore, feeding juveniles on prey eggs and adult females on maize pollen is better for long-term preservation of N. californicus females than continuous feeding on maize pollen due to a shorter developmental duration, a higher egg production and more favourable life table values.     

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.     

Biology of Leipothrix dipsacivagus (Acari: Eriophyidae), a candidate for biological control of invasive teasels (Dipsacus spp.)

The present study describes key aspects of the biology of Leipothrix dipsacivagus, an eriophyid mite that is under study as a biological control candidate of Dipsacus fullonum and D. laciniatus (Dipsacaceae). Preliminary host-specificity tests have shown that it can develop and reproduce only on Dipsacus spp. (teasels). Studies were conducted in a laboratory at 26 ± 2^oC with 16 h of light per day. Mites for the stock colony were collected from D. laciniatus in Klokotnitsa, Bulgaria and reared on rosettes of D. laciniatus in the laboratory. Unfertilized L. dipsacivagus females reared in isolation from the juvenile stage produced male offspring only, while progeny of fertilized females were of both sexes, suggesting arrhenotokous parthenogenesis with haplodiploid sex determination. Experiments were designed to compare male progeny from fertilized females to males from unfertilized females and to compare males and females from fertilized females. Male progeny of virgin mothers had significantly longer durations of active immature stages and total egg-to-adult period than male progeny of fertilized females. Female progeny had significantly longer durations of egg incubation, active immature stages and egg-to-adult period than male progeny from fertilized mothers. Adult longevity was significantly greater in females than in males. Fertilized females produced significantly more eggs per day and overall than virgin females. The results of this study suggest that fertilization status of L. dipsacivagus females can affect both their own fecundity and the development of their male progeny.     

Temperature and sex allocation in a spider mite

Although temperature is the most important environmental factor regulating arthropod development and reproduction, its influence on sex allocation in haplodiploid arthropods remains largely unexplored. We investigated under laboratory conditions how maternal age and temperature mediate offspring sex ratio of the spider mite Tetranychus mcdanieli (Acari: Tetranychidae). Over nine temperature regimes, female-biased sex ratios were consistently observed, varying from 57 to 87% among progeny produced over lifetime. Spider mite sex ratio was affected by maternal age: more male progeny were produced at both the beginning and the end of the female lifespan, yielding a dome shaped curvilinear relationship. This pattern of variation with age probably results from constraints on using sperm at young ages and sperm depletion or viability at older ages. We found a significant curvilinear relationship between temperature and sex ratio. The proportion of female offspring was lowest at intermediate temperatures and highest at extreme temperatures. We suggest that increased female-biased sex ratio at extreme temperatures is an evolutionary response of spider mites to deteriorating habitats as, in the Tetranychidae, females have better capacities than males to disperse and survive under harsh conditions.     

Pair bond duration influences paternal provisioning and the primary sex ratio of brown thornbill broods

Parents should vary their level of investment in sons and daughters in response to the fitness costs and benefits accrued through male and female offspring. I investigated brood sex ratio biases and parental provisioning behaviour in the brown thornbill, Acanthiza pusilla, a sexually dimorphic Australian passserine. Parents delivered more food to male-biased than female-biased broods. However, factors determining parental provisioning rates differed between the sexes. Female provisioning rates were related to brood sex ratio in both natural and experimental broods with manipulated sex ratios. In contrast, male provisioning rates were not affected by brood sex ratio in either natural or experimental broods. However, males in established pairs provisioned at a higher rate than males in new pairs. Data on the sex ratio of 109 broods suggest that female brown thornbills adjust their primary sex ratio in response to pair bond duration. Females in new pairs produced broods with significantly fewer sons than females in established pairs. This pattern would be beneficial to females if the costs of rearing sons were higher for females in new than established pairs. This may be the case since females in new pairs provisioned experimental all-male broods at elevated rates. The condition of nestlings also tended to decline more in these all-male broods than in other experimental broods. This will have additional fitness consequences because nestling mass influences recruitment in thornbills. Female thornbills may therefore obtain significant fitness benefits from adjusting their brood sex ratio in response to the status of their pair bond. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Does variation of sex ratio enhance reproductive success of offspring in tawny owls (Strix aluco)

Tawny owls, Strix aluco, laid female-biased clutches on territories with more abundant prey (field voles) in June, the month that chicks fledge. This appeared to enhance the subsequent reproductive success of fledglings, as in 1995 there was a significant correlation between the number of chicks fledged by adult females and the June vole abundance in the territory on which they were reared as chicks. This relationship did not hold for males. Since tawny owls lay eggs in March, these results indicate that owls are able to predict the June vole numbers on their territory, and respond by producing more of the sex most likely to gain a long-term benefit when resources are good.     

Effect of prey stage on life-table attributes of a genetically manipulated strain ofMetaseiulus occidentalis (Acari: Phytoseiidae)

Metaseiulus occidentalis females from the carbaryl-organophosphate-sulfur resistant strain (COS) lived longer (25.3 days versus 19.7 days), had a higher total fecundity (43.8 eggs female^–1 versus 33.6 eggs female^–1) and a higher daily fecundity rate (2.4 eggs female^–1 day^–1 versus 2.0 eggs female^–1 day^–1), and exhibited a higher intrinsic rate of increase (0.243 individuals female^–1 day^–1 versus 0.182 individuals female^–1 day^–1) and shorter generation time (13.9 days versus 17.0 days), at 24–28°C, 47–56%rh under continuous fluorescent light, when reared on a diet of 0–48-h-old eggs rather than a diet of mixed actives ofTetranychus pacificus McGregor on bean leaf disks. The sexratio of the progeny was female-biased for both diets, 2.1 females to 1 male forM. occidentalis reared on eggs and 2.0:1 : forM. occidentalis reared on mixed actives, suggesting that diet influences sex-ratio in some unknown way.There was no significant difference in oviposition rates for repeatedly-mated and once-matedM. occidentalis females reared on a diet of younger (0–24-h old) eggs compared to a diet of older (72–96-h old) eggs ofT. pacificus.The COS strain ofM. occidentalis exhibited life-table parameters comparable to the other strains reported in the literature, suggesting that the reproductive attributes of this acarine predator were not reduced as a result of artificial laboratory selection. Diet, a biotic factor, produced substantial differences in life-table parameters, suggesting that this factor can influence conclusions regarding the potential efficacy of biological control agents.     

A sex allocation cost to polyandry in a parasitoid wasp

The costs and benefits of polyandry are central to understanding the near-ubiquity of female multiple mating. Here, we present evidence of a novel cost of polyandry: disrupted sex allocation. In Nasonia vitripennis, a species that is monandrous in the wild but engages in polyandry under laboratory culture conditions, sexual harassment during oviposition results in increased production of sons under conditions that favour female-biased sex ratios. In addition, females more likely to re-mate under harassment produce the least female-biased sex ratios, and these females are unable to mitigate this cost by increasing offspring production. Our results therefore argue that polyandry does not serve to mitigate the costs of harassment (convenience polyandry) in Nasonia. Furthermore, because males benefit from female-biased offspring sex ratios, harassment of ovipositing females also creates a novel cost of that harassment for males.     

Effects of founder population size on the performance of Orius laevigatus (Hemiptera: Anthocoridae) colonies

Orius laevigatus (Hemiptera: Anthocoridae) is a key predator of thrips and is mass reared in large numbers for use in biological control. The aim of this study was to evaluate the effect of founder population size on the biological and behavioral performance of O. laevigatus over time. Laboratory lines were started from 1, 10 and 50 founder couples from 750 adults collected in the field and their performance was evaluated at the 5th–6th and 10th–11th generations. Adaptation to the captive rearing situation occurred in the 10 and 50 founder couples lines while it failed in the 1 founder couple line. The intrinsic rate of natural increase (r_m) increased and the period for doubling the population (D) decreased over the generations in the 10 and 50 founder couples lines, while (r_m) decreased and (D) increase in the 1 founder couple line. Also, consumption of Frankliniella occidentalis prey was significantly lower for females from the 1 founder couple line at the 5th generation compared to females from the 10 and 50 founder couples lines. Females of laboratory lines of all founder couples did not respond to odours from thrips infested plants during the 5th and 10th generations, whereas wild females strongly reacted to these odours. We suggest that the lack of reaction to infested plant volatiles may be due to the artificial rearing method where mass reared predators do not experience an infested crop. The results showed that the 1 founder couple line differed from the 10 and 50 founder couples lines, suggesting that bottlenecking had an effect at that level. However, no difference was found between the 10 and 50 founder couples lines which suggest that these founder numbers can be used to start laboratory-reared O. laevigatus lines without a significant loss in quality of its relevant biological characteristics.     

Quantity matters: male sex pheromone signals mate quality in the parasitic wasp Nasonia vitripennis

Sexual selection theory asserts that females are well adapted to sense signals indicating the quality of potential mates. One crucial male quality parameter is functional fertility (i.e. the success of ejaculates in fertilizing eggs). The phenotype-linked fertility hypothesis (PLFH) predicts that functional fertility of males is reflected by phenotypic traits that influence female mate choice. Here, we show for Nasonia vitripennis, a parasitic wasp with haplodiploid sex determination and female-biased sex ratios, that females use olfactory cues to discriminate against sperm-limited males. We found sperm limitation in newly emerged and multiply mated males (seven or more previous matings) as indicated by a higher proportion of sons in the offspring fathered by these males. Sperm limitation correlated with clearly reduced pheromone titres. In behavioural bioassays, females oriented towards higher doses of the synthetic pheromone and were attracted more often to scent marks of males with a full sperm load than to those of sperm-limited males. Our data support the PLFH and suggest that N. vitripennis females are able to decrease the risk of getting constrained to produce suboptimal offspring sex ratios by orienting towards gradients of the male sex pheromone.