Survivorship,potential, and realized fecundity ofBiosteres tryoni [Hymenoptera: Braconidae], a larval parasitoid ofCeratitis capitata [Diptera: Tephritidae]

Laboratory-rearedBiosteres tryoni (Cameron) reached peak ovarian maturation on the 5^th and 7^th days after emergence in unmated and mated ♀♀, respectively. When ♀♀ were not allowed to oviposit, contents of the oocytes were hydrolyzed and reabsorbed. However, when hosts were available for oviposition a significant increase of egg maturation rate was noticed. Mean realized fecundity during the life span was 71.7±8.1 and 69.0±6.7, and mean longevity was 13.5±0.9 and 15.1±0.6 days for mated and unmated ♀♀ respectively.      

Biological studies including population growth parameters of the predatory miteAmblyseius barkeri [Acarina.: Phytoseiidae] at 25°C in the laboratory

Amblyseius barkeri (Hughes) fed onThrips tabaci (Lind.) at 25°C showed an average duration of 2.2, 0.8 and 3.2 days for the egg, larval and nymphal stages, with mortalities at 1.0, 1.0 and 3.1%, respectively. Females represented 63% of the population and required multiple matings for optimal fertility. The oviposition period was 20.3 days and the average oviposition rate 2.3 eggs per day. The intrinsic rate of increase was 0.22 per day. The expected life span was 29.6 days for ♀♀ and 27.4 days for ♂♂.A. barkeri ♂♂ and ♀♀ both consumed 3.3 nymphs of thrips per day (mean value for the feeding stages). The larva does not take up food. In the absence of thripsA. barkeri was able to consume two-spotted spider mites,Tetranychus urticae (Koch), and their eggs, adult broad mites,Polyphagotarsonemus latus (Banks), and pollen of various plants. Cannibalism was observed when food was lacking. Certain morphological features, egglaying, mating and predatory behaviour are described.      

Host selection by the heteronomous hyperparasitoid Encarsia pergandiella: multiple-choice tests using Bemisia argentifolii as primary host

The ovipositional patterns of the heteronomous hyperparasitoid Encarsia pergandiella Howard (Hymenoptera: Aphelinidae) in the presence of its primary host Bemisia argentifolii Bellows & Perring (Hemiptera: Aleyrodidae), and in the presence or absence of conspecific and heterospecific secondary hosts (Encarsia formosa Gahan andEretmocerus mundus Mercet; Hymenoptera: Aphelinidae) were examined to assess host species preferences. Host preferences by heteronomous hyperparasitoids may affect the relative abundance of co-occurring parasitoid species and may influence host population suppression by the parasitoid community. Four combinations of hosts were tested: (1) B. argentifolii, E. mundus, and E. formosa, (2) B. argentifolii, E. formosa, and E. pergandiella, (3) B. argentifolii, E. mundus, and E. pergandiella, and, (4) B. argentifolii, E. mundus, E. formosa, and E. pergandiella. Arrays of hosts (24) were constructed in Petri dishes using leaf disks, each bearing one host. Thirty arrays of each host combination were exposed to single females for 6 h. All hosts were dissected to determine number of eggs per host. Encarsia pergandiella parasitized E. formosa hosts as frequently as E. mundus hosts. However, E. pergandiella parasitized either of these heterospecific hosts more frequently than conspecific hosts in treatments including two secondary host species. When a third parasitoid species was included in host arrays, E. pergandiella parasitized conspecific hosts as frequently as heterospecific hosts. Developmental stage of the hosts did not significantly influence host species selection by E. pergandiella. Our results indicate that host selection and oviposition by heteronomous hyperparasitoids like E. pergandiella, vary with the composition of hosts available for parasitization, and suggest a preference for heterospecific over conspecific secondary hosts.     

The effect of host age ofLymantria dispar larvae [Lep.: Lymantriidae] on the development ofGlyptapanteles liparidis [Hym.: Braconidae]

The endoparasitic development ofG. liparidis was examined in 3 different host stages of gypsy moth larvae. Hatching ofG. liparidis-larvae occurred 3 to 5 days after oviposition in hosts parasitized during their premoulting period, and after 5 to 7 days in those parasitized in the 3^rd midinstar state. The parasites generally moulted to the 2^nd larval instar between the 11^th and 13^th day in the first group, and between the 13^th and 15^th day in the latter, when they had reached a volume of 0.04–0.05 mm³. The positive correlation between host ecdysis and the ecdysis of 1^st stadium larvae to L₂ suggested that host moulting influenced the development of the parasitoid larvae. Emergence from the host larvae occurred at 20°C after 27 days on average, and coincided with the parasites moulting to the 3^rd instar. Five to 7 days after spinning their cocoons near the developmentally arrested host larva, the male, and 1 to 2 days later the female wasps eclosed. Due to the variation in the number of parasites per host, no difference was observed between the hosts parasitized at various stages; however, a tendency for later parasitized hosts to contain more parasite larvae was evident. The nutritional conditions of the moth parental generation influenced both host and parasite development. On the other hand no influence of host age was observed on emergence dates of larvae and wasps.      

Host age and host selection byTrichogramma nubilale

Selection byTrichogramma nubilale (Ertle & Davis) for different aged egg masses ofOstrinia nubilalis (Hübner) was examined by separating components of parasitism. FemaleT. nubilale were exposed to egg masses of 2 different ages (freshly laid egg masses versus 1, 2, 3 or 4 day old egg masses), and the age of the initial egg mass inspected, number and percent of eggs and egg masses parasitized, and the emergence rate of pharate parasitoids was recorded. When there was a large difference in age between young and old egg masses, ♀♀ inspected initially more young egg masses (>2 days difference), parasitized a greater percent of young egg masses (>2 days difference), and parasitized more eggs/parasitized egg mass in young egg masses (>3 days difference) than old egg masses. Females did not discriminate between host age when hosts were similar in age. Successful emergence of parasitoids was greater from younger eggs than older eggs, although emergence was better from one-day-old eggs than from fresh eggs. These results imply that when the hosts were sufficiently different in age, ♀♀ were attracted to younger egg masses more than older egg masses, and they tended to remain ovipositing for longer periods of time on younger egg masses than older egg masses. Moreover, ♀♀ appeared to prefer to oviposit in hosts in which their young are more likely to complete development.      

Bionomics ofTetrastichus sesamiae [Hymenoptera: Eulophidae], a pupal endo-parasitoid ofMaruca testulalis [Lepidoptera: Pyralidae]

The biology and behaviour ofTetrastichus sesamiae Risbec, a pupal endoparasitoid ofMaruca testulalis Geyer, were studied under laboratory conditions. Most adults emerged from the host pupae between 08.00 h and 09.00 h and mating and oviposition started almost immediately. Both ♂♂ and ♀♀ mated repeatedly, and each ♀ could lay eggs for up to 6 days and in up to 5 host pupae. Progeny production ranged from 0–263 offspring per ♀ and adult longevity was from 4.3–13.9 days. The quality of food available to the adults was a major factor influencing progeny production, and longevity. The species was capable of parasitizing and completing development on pupae of such other major Lepidopteran crop pests asChilo partellus Swinhoe,Busseola fusca Fuller,Eldana saccharina Wlk andSpodoptera exempta Wlk, an important finding for biological control of these pests under intercropped agro ecosystems. Apart from parasitism, ♀♀ ofT. sesamiae also caused considerable mortality by stinging, and presumably paralyzing, host pupae without ovipositing.      

Studies of four thelytokous puerto rican isolates ofMuscidifurax uniraptor [Hymenoptera: Pteromalidae]

Field collections in Puerto Rico revealed 4 behaviorally distinct isolates ofMuscidifurax uniraptor Kogan & Legner that differed initially in diapause and nondiapause emergence, and the age when ♀ progeny were produced. Subsequent F₁ and F₂ progeny differed in sex ratio and total progeny production. Mating F₂ ♀ ♀ from nondiapause isolates to naturally emerging ♂ ♂ from thelytokous populations significantly reduced total progeny and the proportion of ♀ ♀ toca. 20 %. These mated ♀ ♀ at first resembled in behavior those which originated from diapausing parents. Random mating within all isolates beginning in the F₃, resulted in a general lower survival and progeny production but was accompanied by a rise in sex ratio toca. 50% ♀ ♀ by the F₆ generation. The interinvolvement of extranuclear and genic factors is considered.      

Progeny production and adult longevity of the mealybug parasitoidsAnagyrus pseudococci,Leptomastix dactylopii,andLeptomastidea abnormis [Hym.: Encyrtidae] in relation to temperature

Progeny production increased and adult longevity decreased with rising temperature within the range 18°C to 30°C for the 3 mealybug parasitoidsAnagyrus pseudococci (Girault),Leptomastix dactylopii Howard andLeptomastidea abnormis (Girault). The Weibull distribution gave a good fit to survival curves for the 3 parasitoids and statistical comparison of Weibullb andc parameters at different temperatures allowed changes in the scale and shape of the curves to be detected. In general, ♀♀ lived longer than ♂♂ for all 3 species, except at high temperature. FemaleL. abnormis attained their maximum progeny production at 24°C and maintained this level up to 34°C. They lived longer than the other 2 parasitoid species at 30°C and showed a type I survival curve throuhout the range of temperatures examined.A. pseudococci andL. dactylopii both required high temperatures (30°C) to attain their maximal progeny production, but werepseudococci tended towards type II, with a larger proportion of the population dying within the first few days.L. dactylopii lived longest at 26°C, with ♀♀ showing a type I survival curve at all temperatures and ♂ survival curves changing from type I to type II at 30°C. The implications of these findings for the population dynamics of the different parasitoids are briefly discussed.      

Horizontal transmission of mycotic infection in adult tsetse,Glossina morsitans morsitans

AdultGlossina morsitans morsitans exposed to wet conidia ofBeauveria bassiana andMetarhizium anisopliae suffered high mortalities ranging from 90 to 100% by 2 weeks post-exposure. Infected ♂ ♂ maintained in the same cages with non-infected ♀♀ throughout the experimental period transmitted the fungal infection to the ♀♀ resulting in mortalities of 65% withB. bassiana and 55% withM. anisopliae. Likewise, infected ♀♀ maintained together with non-infected ♂♂ transmitted the infection to the ♂♂ resulting in mortalities of 75% withB. bassiana and 45% withM. anisopliae. Female tsetse flies infected withB. bassiana andM. anisopliae and maintained in the same cages with non-infected ♀♀ also transmitted infection to the non-infected tsetse resulting in mortalities of 62% and 48% withB. bassiana andM. anisopliae respectively. Infected tsetse exposed to non-infected tsetse of the opposite sex for only 30 min were also able to transmit the fungal infection. Pupae produced by female tsetse infected withB. bassiana andM. anisopliae exhibited higher pupal mortality than those produced by non-infected ♀♀. However, pupae exposed directly to dry spores ofB. bassiana andM. anisopliae had no increase in pupal mortality but adults emerging from theB. bassiana-exposed pupae had markedly reduced longevity.      

Larvicidal activity of lectins onLucilia cuprina: mechanism of action

Foraging behaviour and host-instar preference of young and old females of the solitary aphid parasitoid,Lysiphlebus cardui Marshall (Hymenoptera: Aphidiidae), were studied in the laboratory. The analysis of interactions between parasitoids and different stages ofAphis fabae cirsiiacanthoidis Scop. (Homoptera: Aphididae) revealed that encounter rates between aphids and parasitoid females and defence reactions of the aphids influenced the degree to which a particular aphid age class is parasitized. Encounter rates between hosts and parasitoid females depended on the foraging pattern of the parasitoid, which varied with age. In mixed aphid colonies patch residence time increased with parasitoid age. Furthermore, younger parasitoids (≦1 day old) laid more eggs into second and third instars, while older parasitoids (≧4 days old) did not show distinct host instar preferences. It is suggested that the oviposition behaviour ofL. cardui is influenced by the physiological state, i.e. the age of the wasp.     

Influence of the egg parasitoidPlatytelenomus busseolae [Hym.: Scelionidae] on the population ofSesamia nonagrioides [Lep.: Noctuidae] in central greece

The egg parasitoid,Platytelenomus busseolae (Gahan) (Hymenoptera: Scelionidae) was recorded from egg masses of the corn stalk borer,Sesamia nonagrioides (Lefebvre) (Lepidoptera: Noctuidae) on maize in the area of Istiaea, Central Greece. Natural parasitism was studied during the years 1986, 1987. A total of 3,382 egg masses, containing 205,227 eggs, was collected from corn fields. Parasitized eggs were found from end July to mid-October. In 1986. 76.2% of the egg masses and 42.8% of all eggs were parasitized, while in 1987, respective values were 27.6% and 12.9%. The inferior performance of the parasitoid in 1987 may be due to the unfavorable winter and summer conditions. Parasitism decreased when the host population was at its peak, but it increased again within 1 to 3 weeks. A small percentage of egg masses was completely parasitized (9.5% and 4.7% for the 2 years respectively). In 1987, the sex ratio, ♀♀/♂♂ was 1.5: 1 and 26.2% of the egg masses produced only ♂♂. Mated ♀♀ produced 3 times more ♀♀ than ♂♂. It seems that the parasitoid, under favorable weather conditions, can play a significant role in the control of the corn stalk borer.      

Oviposition pattern byCotesia ayerzai [Hym.: Braconidae] onAscia monuste [Lep.: Pieridae] under laboratory conditions

Various aspects of the parasitims of caterpillars ofAscia monuste orseis byCotesia ayerzai, were studied in laboratory choice tests. Individual ♂♂ were found to be extremely variable in ovipositional duration, as well as in the number of eggs oviposited. To simulate parasitoid dispersal, or low host density, we isolated ♂♂ for 60 min following initial exposure and compared our tests with ♂♂ which had not been isolated. No differences in (1) the number of eggs oviposited per host instar; (2) the conditional probability of host encounter; (3) host acceptance; and (4) the conditional probability of instar specific parasitism were found between these groups. Held ♂♂ did, however, reject hosts after ovipositor insertions in a higher proportion.      

Influence of previous herbivory on behavior and development of Spodoptera exigua larvae on glanded and glandless cotton

The solitary endoparasitoid Aenasius vexans Kerrich (Hymenoptera: Encyrtidae) is used for augmentative releases against the cassava mealybug, Phenacoccus herreni Cox & Williams (Sternorrhyncha: Pseudococcidae), an important pest on cassava in South America. In light of the need for large numbers of high quality females, experiments were conducted on host stage suitability and sex allocation. In choice and no-choice experiments, individual female wasps were offered second and third instar, as well as adult, hosts. During the first five days after emergence, the wasps showed a steady increase in the number of hosts they successfully parasitised per day, but the respective secondary sex ratio for each instar remained constant. Parasitism was highest for third instar hosts in no-choice tests, while in choice tests parasitism was highest in both third instars and adults. The later the developmental stage of the host at oviposition, the faster the parasitoids developed and emerged, and for each host stage, the development time of males was shorter than for females. The sex ratio of the wasps emerging from hosts that were parasitised as second instars was strongly male-biased, while the apparently preferred later stages yielded significantly more females than males. Female and male A. vexans emerging from hosts parasitised at the third instar were significantly larger than for the other stages. This may explain the preference for the third instar as well as the female-biased sex ratio, as size is usually positively correlated with higher fitness, especially in females. The results suggest that third instar hosts are the most suitable for rearing high numbers of large females.     

Does superparasitism improve host suitability for parasitoid development? A case study in the Microplitis rufiventris-Spodoptera littoralis system

Superparasitism refers to the oviposition behavior of parasitoid females who lay their eggs in an already parasitized host. Recent studies have shown that allocation of additional eggs to an already parasitized host may be beneficial under certain conditions. In the present work, mortality of Microplitis rufiventris wasps was significantly influenced by both host instar of Spodoptera littoralis larvae at parasitism and level of parasitism. In single parasitization, all host instars (first through sixth) were not equally suitable. Percentage of emergence success of wasp larvae was very high in parasitized first through third (highly suitable hosts), fell to 60% in the fourth instar (moderate suitable) and sharply decreased in the penultimate (5th) instars (marginally suitable). Singly parasitized sixth (last) instar hosts produced no wasp larvae (entirely unsuitable), pupated and eclosed to apparently normal adult moths. The scenario was different under superparasitism, whereas supernumerary individuals in the highly suitable hosts were almost always killed as first instars, superparasitization in unsuitable hosts (4th through 6th) had significant increase in number of emergence success of wasp larvae. Also, significantly greater number of parasitoid larvae successfully developed in unsuitable hosts containing three wasp eggs than counterparts containing two wasp eggs. Moreover, the development of surplus wasp larvae was siblicidal in earlier instars and nonsiblicidal gregarious one in the penultimate and last “sixth” instars. It is suggested that the optimal way for M. rufiventris to deal with high quality hosts (early instars) is to lay a single egg, while the optimal way to deal with low quality hosts (late instars) might be to superparasitize these hosts.     

Inheritance of dicrotophos resistance in greenhouse whitefly

We studied inheritance of resistance to dicrotophos in greenhouse whitefly,Trialeurodes vaporariorum Westwood (Homoptera, Aleyrodidae). Compared with females from a field-collected susceptible strains (S), females from a resistant strain (R) ofT. vaporariorum derived from heavily treated cotton fields had a 28-fold greater LC₅₀ to dicrotophos in laboratory bioassays. Concentration-mortality lines obtained from female progeny of reciprocal F₁ crosses (R_♀ XS_♂ and S_♀ XR_♂) were similar, suggesting that inheritance of dicrotophos resistance was autosomal and not influenced by maternal effects. Responses of F₁ female progeny were similar to those of the parental S strain, indicating that the resistance was partially recessive (degree of dominance, D, was −0.61). Mortality observed in female progeny obtained from a backcross (F_1♀ XR_♂) corresponded more closely to expectations derived from polygenic models than to expectations from a monogenic model. The estimated number of effective factors (sensu Lande, 1981) contributing equally to resistance was three.     

Host selection and survival of the parasitoidEncarsia formosa on greenhouse whitefly,Trialeurodes vaporariorum, in the presence of hosts infected with the fungusAschersonia aleyrodis

Successful control of greenhouse whitefly may be achieved by complementary activity of the parasitoidEncarsia formosa and the fungusAschersonia aleyrodis. One way to obtain an additive mortality effect of both entomopathogen and parasitoid would be achieved by the selection of healthy hosts by the parasitoid and rejection of fungus-infected hosts. Third and fourth instar larvae ofTrialeurodes vaporariorum which had been treated with a spore suspension ofA. aleyrodis 0, 4, 7, 10 or 14 days beforehand, were presented to female parasitoids. The parasitoids adopted the oviposition posture on untreated hosts as well as on treated hosts, irrespective of the different stages of infection in the hosts. However, significantly more hosts were parasitized byE. formosa in the control treatment than in the fungal treatment. The parasitoids offered treated hosts, showed rejection behaviour after probing on hosts showing detectable signs of infection (containing hyphal bodies or mycelium in the haemolymph). For instance, when hosts were offered seven days after spore treatment, the parasitoids showed an oviposition posture on a total of 83 (95.4%) out of 87 infected larvae, but laid only 4 eggs (4.6%). In contrast, on 48 (94.1%) out of 51 noninfected (or showing no detectable signs of infection) hosts an oviposition posture was adopted and 40 eggs (78.4%) were found after dissection. When infected hosts were encountered the oviposition posture lasted less than 1′40″ while rejection of non-infected hosts occurred after more than 1′40″. Other experiments were carried out offering treated hosts for 24 h to the parasitoids. The hosts were dissected afterwards. Again, significantly more eggs were laid in the non-infected hosts. When hosts were parasitized shortly after fungal spore treatment they were colonized by the fungus and the parasitoids did not develop. Transmission of the entomopathogen after probing infected hosts was observed to a limited extent. In conclusion,A. aleyrodis andE. formosa can be used together in a glasshouse situation. The parasitoid will be most effective when introduced more than seven days after application ofA. aleyrodis, because from that time onwards it is able to detect and reject fungus-infected hosts.     

Selection on the male hemizygous genotype in arrhenotokous insects and mites

Arrhenotokously reproducing Hymenoptera and Acarina include many important natural enemies. This reproductive system offers the opportunity of selection on hemizygous (♂ ♂), with the attendant advantages of an unmasked genotype fully exposed to selection, in creased frequency of genotypes expressing rare genes, and enhanced discrimination of characters, in all-♂ populations produced by virgin (♀ ♀). Increased selection intensity and reduced genetic drift may offer additional advantages. The method is limited to characters displayed by (♂ ♂), and may require labor-intensive techniques and species-specific research. The method has been shown to be practicable withAphytis holoxanthus DeBach (Hymenoptera: Aphelinidae), an important parasite of the Florida red scale,Chrysomphalus aonidum (L.) (Homoptera: Diaspididae).      

Egg Size, Intrinsic Competition, and Lethal Interference in the Parasitoids Encarsia pergandiella and Encarsia formosa

Recent population dynamic theory predicts that disruption of biological control may occur when one parasitoid species' superiority in intrinsic competition is associated with a lower ability to find and exploit hosts (i.e., ability in extrinsic competition). One might expect such a trade-off, for instance, if parasitoids with larger (and fewer) eggs are more likely to prevail in intrinsic competition than species with smaller (and more numerous) eggs. We tested the idea that relative egg size could be used to predict the outcome of intrinsic competition in two closely related endoparasitoids, Encarsia pergandiella Howard and Encarsia formosa Gahan. Contrary to expectation, the parasitoid species with smaller eggs, E. pergandiella, prevailed in intrinsic competition, regardless of the order that hosts were exposed to the two species. In a literature survey, we found four studies of competing pairs of endoparasitoid species for which: (a) egg size estimates were available and (b) one species was consistently superior in intrinsic competition. In three of the four studies, the small-egged species prevailed in intrinsic competition, as we also found. Although E. formosa lost in intrinsic competition, this species negatively affected E. pergandiella's progeny production by host feeding on and killing hosts containing E. pergandiella eggs. E. formosa females also host fed on conspecific-parasitized hosts. As a mechanism of both intra- and interspecific interference competition, host feeding on parasitized hosts contradicts assumptions about the nature of interference competition in existing population dynamics models.     

Paternal influence in males ofMuscidifurax raptorellus [Hymenoptera: Pteromalidae]

Hybrid ♀♀ from cohorts of solitary and gregarious populations ofMuscidifurax raptorellus harboring one-half the genome of either parent, are accompanied in emergence by ♂♂ most typical of the original female parent used to create the hybrid. However, these ♂♂ also show small influences of their mother's mating partner. This suggests the existence of a extranuclear phase to the inheritance of gregarious oviposition in this species, and biparental influences in the hymenopteran male.

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Résumé Les ♀♀ hybrides de cohortes de populations solitaires et grégaires deMuscidifurax raptorellus extériorisant la moitié du génome de chaque parent sont accompagnées à l'émergence par des ♂♂ les plus typiques de la femelle d'origine utilisée pour réaliser l'hybride. Cependant ces ♂♂ manifestent aussi de légères influences du partenaire de leur mère. Ceci suggère l'existence d'une phase extranucléaire à l'héritage d'une ponte grégaire dans cette espèce et des influences biparentales chez le male d'hyménoptères.

     

Effects of host instar and size on parasitization efficiency and life history parameters ofOpius dissitus

The braconid parasitoidOpius dissitus Muesebeck (Hymenoptera, Braconidae) produced 1.7 to 3 times more offspring when provided second and third instar leafminers (Liriomyza sativae Blanchard) as compared to first instars. Females arising from parasitization of different instars did not differ significantly in numbers of chorionated eggs in their ovaries at adult eclosion. Development time was prolonged by about two days when parasitoid oviposition occurred in first, as compared to third instar hosts. Parasitoid length was positively correlated with host weight (r²=0.75). Because only 7% of variation in host weight could be explained by host density, parasitoid length varied considerably among hosts reared at the same density. Longevity and lifetime fecundity of parasitoids were inversely related to the weight of their hosts.