Factor Affecting Sex Ratios of a Mermithid Parasite of Mosquitoes

The ratio of male to female Reesimermis nielseni Tsai and Grundmann, a nematode parasite of mosquito larvae, increased as the number of parasites per host increased. Hosts with a single nematode produced 9% males compared with essentially 100% males in hosts with more than 7 parasites; hosts with 3 nematodes produced about equal numbers of males and females. Males of R. nielseni generally emerged before females because of the earlier death of multiple-infected mosquitoes. The species of the host mosquito influenced the sex ratio, but the size of a specific host at the time of invasion did not. Host diet also had a noticeable influence on the sex ratio of the nematode: singly infected hosts from a starved population produced 92% males compared with 13% in the normally fed group. The importance of these factors in the mass rearing of R. nielseni is discussed.     

Effects of male-female rations on mating and egg production in Octomyomermis muspratti (Mermithidae: Nematoda)

Octomyomermis muspratti, a mermithid parasite of mosquito larvae, molted to the adult stage in 4–16 days. When male-female ratios were relatively high (1:2), mating and oviposition began soon after maturation, and oviposition was completed in 21–40 days. The number of females developing eggs increased with a corresponding increase in the male-female ratio. About 60% of the females completed oviposition at male-female ratios of 1:20, and about 100% completed oviposition as the male-female ratios were increased to 1:2. No egg development was observed in the absence of males. Male O. muspratti were capable of mating with a number of females and remained reproductively active for at least 50 days after a maturation at 26–28°C.     

Effects of Host Diet on Romanomermis culicivorax,a Mermithid Parasite of Mosquitoes

When larval mosquitoes (Aedes aegypti) infected with the mermithid nematode Romanomermis culicivorax were fed on a diet low in quantity or protein content or both, the number of postparasites which emerged from the hosts decreased and host mortality increased marginally. Parasitic development was prolonged and became asynchronous in nutritionally deprived hosts. Nematodes emerged from insects infected by more than one nematode before the remaining juveniles comprising such infections had completed parasitic development; this resulted in substantial reductions in postparasite nttmbers. Host development was retarded by low protein and/or reduced diets. Postparasites emerging from second and third instars were reduced in size and in the amount of stored nutriment compared to those recovered from hosts fed on a high protein diet ad libitum. A greater proportion of the rnermithids developed into males in hosts fed on reduced diets but not in hosts fed on low protein diets.     

Penetration and Development of the Mermithid Nematode Reesimermis nielseni in Eighteen Species of Mosquitoes

The susceptibility of 18 species of mosquitoes to the infective stage of the mermithid nematode Reesimermis nielseni was compared to that of Culex pipiens quinquefasciatus. Thirteen species were more susceptible, with three Anopheles species and Culiseta inornata the most susceptible. Aedes triseriatus, Culex territans, and Psorophora ferox were highly resistant. Resistance to R. nielseni appeared to be behavioral, physical, or physiological, and some host species exhibited one or more types of defense mechanisms. No noticeable differences were apparent in the degree of susceptibility of a native-and a laboratory strain of C. p. quinquqfasciatus to R. nielseni.     

Evidence for Strategic Sex Allocation in the Guppy (Poecilia reticulata): Brood Sex Ratio and Size Predict Subsequent Adult Male Mating Success

Sex allocation theory predicts that females should produce more sons when the reproductive success of sons is expected to be high, whereas they should produce more daughters, not daughters when the reproductive success of sons is expected to be low. The guppy (Poecilia reticulata) is a live‐bearing fish, and female guppies are known to produce broods with biased sex ratios. In this study, we examined the relationship between brood sex ratio and reproductive success of sons and daughters, to determine whether female guppies benefit from producing broods with biased sex ratios. We found that sons in male‐biased broods had greater mating success at maturity than sons in female‐biased broods when brood sizes were larger. On the other hand, the reproductive output of daughters was not significantly affected by brood sizes and sex ratios. Our results suggest that female guppies benefit from producing large, male‐biased brood when the reproductive success of sons is expected to be high.     

Effects of host larval stage preferences and diet on life history traits of Diadegma mollipla,an African parasitoid of the Diamondback Moth

Knowledge of the biological attributes of parasitoids plays a fundamental role in improved management of their pest hosts. Diadegma mollipla is a major indigenous parasitoid of the Diamondback Moth (DBM), Plutella xylostella (Linnaeus), in Eastern Africa. Two laboratory experiments were conducted to determine the DBM larval stage preferences of D. mollipla and their influence on life history traits and the effect of different diets (20% honey, 20% sugar solution, water or nothing) on adult parasitoid longevity. In the first study, 60 DBM larvae (15 each of larval stages L1–L4) were exposed to either one adult parasitoid female or four competing females in choice experiments. When DBM larvae were exposed to one parasitoid, in terms of parasitism, L1 and L2 were found to be the most preferred stages, followed by L3 and then L4. Sex ratio (females/males) increased with host larval stage from 1.3 to 3.1. Pre-imaginal development time was longer in L1 than in the other three stages. When exposed to four parasitoids, no significant differences in parasitism rates and progeny production were obtained between DBM larval stages. However, sex ratio increased from 0.8 in L1 to 4.1 in L4. Developmental time was longest in L4, intermediate in L1 and L3, and shortest in L2. In the diet experiment, honey and sugar were found to significantly increase longevity for D. mollipla adults of both sexes. Overall, combined adult male and female longevity was 24.5, 9.9, 2.2 and 1.9 days when fed on honey, sugar, water or nothing, respectively.     

Effect of the Entomogenous Nematode Nemplectana carpocapsae on the Tachinid Parasite Compsilura concinnata (Diptera: Tachinidae)

The entomogenous nematode Neoaplectana carpocapsae and its associated bacterium, Xenorhabdus nematophilus, could not infect the pupal stage of the tachinid Compsilura concinnata through the puparium. N. carpocapsae had an adverse effect on 1-, 2- and 3-day-old C. concinnata larvae within the armyworm host in petri dish tests. All 1-day-old larvae treated with nematodes died in their hosts, whereas 61% and 69% of 2- and 3-day-old larvae treated with nematodes, respectively, died. However, the survivors developed to adults. Nine to thirty-seven percent of adult tachinids which emerged from nematode-treated soil (50 nematodes/cm²) were infected with N. carpocapsae. The nematode adversely affects C. concinnata directly by the frank infection of the tachinid and indirectly by causing the premature death of the host which results in tachinid death.     

Laboratory Culture and Life History of Heleidomermis magnapapula in Its Host, Culicoides variipennis (Diptera: Ceratopogonidae)

The mermithid parasite Heleidomermis magnapapula was maintained in larvae of the midge Culicoides variipennis for 20 months in enamel pans containing nutrient-rich water and polyester pads as a substrate. Inseminated female mermithids were introduced to the pad surface when the host was in the late second or early third-instar. Host larvae were harvested from the pans 9 days after exposure and held in tap water for nematode emergence. Preparasite yield was positively correlated with female nematode size and averaged 1,267 preparasites/female. Male and female nematodes emerged an average of 12.2 and 13.4 days after host exposure, respectively. Supplemental host food (Panagrellus) during the final days of parasitism did not alter time of emergence. Parasites emerging singly were 64% females, whereas superparasitized hosts yielded males (up to nine/host). Nematode carryover into the adult midge normally occurred at a level of 0.5-2.5%. Parasite load (nematodes/ parasitized individual) in midge adults was lower than that of larvae from the same cohort, and adult midges were more likely to harbor female parasites. Exposure of fourth-instar host larvae resulted in higher levels of adult parasitism (up to 17%).     

A Comparative Study of Sex Ratio and Clutch Size in Gregarious Ichneumonoid Wasps

Some convincing support for sex ratio theory comes from the cross-species relationship between sex ratio and brood size in gregarious bethylid wasps (Hymenoptera: Bethylidae), in which the proportion males declines as brood size increases as predicted under local mate competition. It is unknown how widely such relationships hold within parasitoid wasps as a whole. We assemble a dataset on sex ratio and brood size for gregarious Braconidae and Ichneumonidae. Their sex ratios deviate substantially from those of bethylids; sex ratios differ widely across species; and they are not significantly related to brood size across species. Several factors explain the heterogeneity in sex ratios including across-species differences in mating system, sex determining mechanism, and sexual asymmetries in larval competition and polyembryony leading to single-sexed broods.     

Superparasitism and sex ratio in the solitary parasitoidEpidinocarsis lopezi

Highly variable sex ratios are found in the solitary parasitoidEpidinocarsis lopezi, both in the field and in a mass-rearing situation. Superparasitism is one of a number of factors which can influence sex ratios in parasitoid Hymenoptera. We show that superparasitism inE. lopezi is common in the field. Sex allocation decisions when parasitizing unparasitized hosts are not different from those with parasitized hosts; neither does differential mortality occur between the sexes in superparasitized hosts. Therefore superparasitism does not contribute to the variable sex ratio ofE. lopezi. Both the occurrence of superparasitism and the sex produced when ovipositing are shown to be functional forE. lopezi.     

Temperature Effects on the Parasitic Phase of Romanomermis culicivorax in Culex pipiens

The developmental period for the parasitic stage of Romanomermis culicivorax in Culex pipiens was determined at constant and fluctuating temperatures. The median developmental times at 15, 18, 20, 27, and 32 C were 27.6, 17.2, 11.5, 7.1, and 5.8 days, respectively. The optimum temperature range for development of the parasitic stage in C. pipiens was 20 to 32 C. The threshold for development was calculated as 10.4 C, and the heat units required for development were 122.2 ± 3.6 day-degrees. Development at fluctuating temperatures conformed to that predicted by the constant temperature data.     

Hexamermis cathetospiculae n. sp. (Mermithidae: Nematoda), a Parasite of the Rice Stemborer,Tryporyza incertulas (Wlk.) (Pyralidae: Lepidoptera) in Malaysia

Hexamermis cathetospiculae n. sp. (Mermithidae: Nematoda), a parasite of the rice stemborer, Tryporyza incertulas (Wlk.), is described. This mermithid was observed to control the stemborer in Malaysia. It is distinguished from other members of the genus by the shape of the vagina, the nearly straight spicules, and the form of the amphids.     

Effects of Selected Insecticides and Nematicides on the In Vitro Development of the Entomogenous Nematode Neoaplectana carpocapsae

The effects of organophosphates (mevinphos, phenamiphos, trichlorfon), carbamates (carbofuran, methomyl, oxamyl), a formamidine (chlordimeform), a synthetic pyrethroid (fenvalerate), a chlorinated hydrocarbon (methoxychlor). and an insect growth regulator (diflubenzuron) on in vitro development and reproduction of Neoaplectana carflocapsae were tested by incorporating each chemical into a nematode rearing medium. Organophosphates and carbamates adversely affected development and reproduction at concentrations ≥ 0.1 mg/ml. Phenamiphos was the most toxic, with no nematode reproduction at 0.01 mg/ml. Inoculated infective juveniles developed to adults with some of the organophosphates and carbamates, but limited or no reproduction occurred. Chlordimeform inhibited development at 1.0 mg/ml, while diflubenzuron, fenvalerate, and methoxychlor did not significantly (P > 0.05) reduced reproduction at 1.0 mg/ml. The organophosphate and carbamate nematicides in use for control of plant-parasitic nematodes may be toxic to N. carpocapsae in the soil.     

Influence of Host Nutrients on the Parasitic Development of Mermis nigrescens (Mermithidae)

Schistocerca gregaria nymphs and adults of both sexes were infected with eggs of Mermis nigrescens. Mermithid larvae grew more slowly in nymphal hosts, and emerging larvae were smaller than those from adult hosts. The longer the larvae remained in the host, the greater their size. Those developing in adult female hosts were longest. Single mermithid larvae that were transferred to a second host continued to grow and were significantly longer at emergence than larvae that developed solely in one host. In adult hosts that were infected with 40-300 M. nigrescens eggs, the percentage of mermithids that became males was strongly dependent on host weight at infective doses of 90 eggs or more. Results are discussed in relation to nutrient stress on the larvae and its importance in developing in vitro culture techniques.     

Effect of Aedes aegypti (Diptera:Culicidae) Age on Sex Ratios in Romanomermis culicivorax (Nematoda:Mermithidae)

First, second, third, early and late fourth-instar larvae, and pupae of Aedes aegypti were infected with Romanomerrais culicivorax and reared at 20, 25, and 30 C. An increase in the ratio of male to female nematodes was observed with increase in host age at the time of infection at each temperature. The number of pupal and late fourth-instar infections was low, but R. culicivorax continued to develop in adult A. aegypti. Since male nematodes were recovered from both male and female hosts infected as late fourth instars or pupae, the sex of the host did not influence the sex of the nematode.     

Effects of conservation strip and crop type on natural enemies of Delia radicum

This 3‐year field study investigated the combined effect of floral resources and perennial shelter habitats (i.e. conservation strips), and crop rotation in supporting natural enemies of the cabbages root fly, Delia radicum. Habitat manipulation with conservation strips increased the overall catches of hymenopteran parasitoids. However, conservation strips did not increase parasitism by either of the two dominant parasitoid species, Trybliographa rapae and Aleochara bipustulata, in any study year. In fact, higher parasitism was found in control plots in the second year. This could be explained by parasitoid mobility and higher patch detectability, as more plants in the control plots were infested with D. radicum larvae. Conservation strips did not result in increased predation of D. radicum eggs. However, the activity densities of two Bembidion species were correlated with egg predation. The species assemblage distribution of epigeal predators was best explained by seasonal period, followed by year and, to a low extent, crop type, while treatment with conservation strips had no effect. However, during the egg laying peak of D. radicum, a higher number of A. bipustulata, an important larval predator was observed in conservation strips during one study year. In conclusion, positive effects of conservation strips were demonstrated for abundance of some natural enemies of D. radicum, but a consistent increase in performance could not be adequately demonstrated due to experimental set‐up, the short timescale and the complex landscape in which our study site was located.     

Temperature Effects on Survival and Development of Heleidomermis magnapapula in the Laboratory

The mermithid Heleidomermis magnapapula Poinar and Mullens, a parasite of the biting midge Culicoides variipennis (Coquillett), was exposed to constant temperatures in the laboratory. Survival of the free-living stages and development times of eggs and the parasitic phase were inversely related to temperature. Average preparasite longevity was 70, 46, 42, and 22 hours at 15.6, 21.1, 26.7, and 32.2 C, respectively. Females survived significantly longer than males. Longevity in days (females/males) at different temperatures was 17.3/11.0 at 4.4 C, 9.0/8.2 at 15.6 C, 5.9/5,1 at 21.1 C, 5.2/4.7 at 26.7 C, and 4.4/3.6 at 32.2 C. Embryogenesis required 44 ± 2 degree days above a thermal minimum of 10.1 C, while parasitic development in host larvae required 214 ± 10 degree days above a thermal minimum of 8.9 C. Parasite responses to temperature were very closely related to temperature-dependent host development patterns.     

Variable Female Mating Positions and Offspring Sex Ratio in the Spider Pityohyphantes phrygianus (Araneae: Linyphiidae)

Chromosomal sex determination and male heterogamety have been thought to seriously impede direct sex ratio control. However, in Pityohyphantes phrygianus, a solitary sheetweb spider with a skewed sex ratio, earlier experimental studies suggested that there are options for female control of offspring sex ratio, if females change their position during the normal mating sequence. Here we show that under natural conditions there is considerable between-female variation in positions, especially after termination of mating. Computer simulations of the orientation of female inner genitalia suggest that sperm are placed in different storage sites depending on the positions adopted. This means that a specific position after mating might potentially influence offspring sex ratio. The variance in offspring sex ratio among females in earlier experiments was binomially distributed, which leads us to conclude that females control the mean sex ratio but do not exercise direct control of the sex of individual offspring.     

Host exploitation and contest behavior in a generalist parasitoid partially reflect quality of distinct host species

Information use determines parasitoid adaptive behavior in general, and host specialization or fitness in specific. Information regarding host suitability could affect sex allocation behavior, host exploitation, or aggressiveness in dyadic contests. In this paper, we relate aggressiveness of the pupal parasitoid Pachycrepoideus vindemmiae (Hymenoptera: Pteromalidae) with sex allocation and host exploitation when presented with different host species. More specifically, we presented parasitoids with puparia of five different Dipteran species: Drosophila melanogaster (Drosophilidae), Musca domestica (Muscidae), Anastrepha obliqua, Anastrepha fraterculus, and Ceratitis capitata (Tephritidae). Puparia of above species greatly varied in size and volume, were parasitized to differing extent and had varying fitness implications for P. vindemmiae. Using a composite measure of selected fitness currencies (i.e., parasitism level, offspring size, longevity and sex ratio), we typified D. melanogaster and A. obliqua as ‘low quality’ hosts for P. vindemmiae while puparia of C. capitata and A. fraterculus were considered of ‘high quality’. In contest dyads, female aggressiveness and host exploitation behavior differed between host species. Wasps exhibited highest frequencies of antennal striking and rival pursuit, and high degrees of puparium mounting, antennating and probing on C. capitata. Antennal striking frequency however was equally high on ‘low quality’ hosts such as D. melanogaster and A. obliqua. This work shows that a generalist parasitoid such as P. vindemmiae assesses host quality when confronted with hosts of differing species, size or nutritional suitability and employs such to define sex allocation, host exploitation, and contest behavior. However, contest and exploitation behavior only partially indicate host quality and broader parasitoid fitness implications. This work has further implications for parasitoid mass rearing and use of P. vindemmiae for biological control of Dipteran pests.