Biology of Tetrastichus howardi (Olliff) (Hymenoptera: Eulophidae): A facultative Hyperparasitoid of stem borers

The biology of the parasitoid Tetrastichus howardi, which was introduced from the Philippines into South Africa for the biological control of the stem borer Chilo partellus, was studied in the laboratory. On average, a T. howardi female produced 101 progeny (92% females) which developed in 4.3 hosts. The upper threshold temperature for development was around 33°C, the minimum threshold was calculated as 12.8°C and the thermal constant as 239.8 day‐degrees. With hosts, females lived 23.6 and males 16.7 days. T. howardi was found to be a facultative hyperparasitoid with a very wide host range. As a primary parasitoid, it developed in pupae of various Lepidoptera as well as Coleoptera and Hymenoptera and as a hyperparasitoid in dipteran and hymenopteran parasitoids. A linear relation was found between the number of emergent parasitoids and the volume of the host from which they emerged. Preference tests indicated that T. howardi preferred to parasitize the phytophagous insects C. partellus and Heliothis armigera, over their parasitoids Xanthopimpla stemmator and Palexorista laxa. The host selection of inexperienced T. howardi females was different from the experienced parasitoids. When given a choice, females showed preference for the host species they were allowed to parasitize previously and the host species from which they had been reared. The use of facultative hyperparasitoids in biological control in general and against stem borers in South Africa in particular is discussed.     

Development and morphological characterization of the immature stages of Tetrastichus giffardianus Silvestri (Hymenoptera: Eulophidae)

The gregarious endoparasitoid Tetrastichus giffardianus Silvestri (Hymenoptera: Eulophidae) is a natural enemy of fruit flies. This parasitoid was previously used to successfully control Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in Hawaii, USA. Despite its importance in the control of fruit fly pests, little is known about the development or characteristics of its preimaginal stages. The aim of this study was to observe the development and morphologically characterize the immature stages of Tetrastichus giffardianus. Tetrastichus giffardianus individuals were reared on C. capitata larvae/pupae under laboratory conditions at a temperature of 25 ± 2 °C, relative humidity of 60 ± 10%, and 12-h photophase. Third-instar C. capitata larvae were exposed to parasitism for 24 h. After parasitism, the pupae were dissected every 24 h to evaluate the stage of development attained by T. giffardianus, and to record their morphological characteristics. A stereomicroscope was used to observe all the immature stages of T. giffardianus. The complete development of T. giffardianus under these conditions was completed within 14 days as follows: egg (duration ? 1 day); first (? 1 day), second (? 1 day), and third (? 2 days) larval instars; pre-pupa (? 2 days); and pupa (? 7 days). The immature stages of T. giffardianus differed sufficiently in their shape, color, and size to allow morphological characterization.     

Ultrastructure of the antennal sensilla of the cockroach-egg parasitoid,Tetrastichus hagenowii (Hymenoptera: Eulophidae)

Eggs of a number of cockroach species are parasitized by Tetrastichus hagenowii. The ultrastructure of the sensilla on the antennae of females and males was examined by scanning and transmission electron microscopy. The females have two types of multiporous plate sensilla while the males have only one. Type 1 is found in females and males and has a relatively thin cuticular wall and many pores, while type 2 is found only in females and has a relatively thick cuticular wall and few pores. Both sexes have nonporous, thick-walled, socketed hairs; multiporous, nonsocketed hairs; multiporous, thick-walled pegs; and terminal hairs. In addition, males have multiporous, nonsocketed, long hairs. The sensilla are similar, in many respects, to the sensilla of other chalcid parasitoids. The antennal sensilla of female T. hagenowii are probably involved in ovipositional behavior. The multiporous, long hairs of the male possibly receive stimuli during mating behavior A chemoreceptive function is proposed for the multiporous plate sensilla.     

Interactions between Spathius agrili (Hymenoptera: Braconidae) and Tetrastichus planipennisi (Hymenoptera: Eulophidae), larval parasitoids of Agrilus planipennis (Coleoptera: Buprestidae)

Three hymenopteran parasitoids native to China are being released in the United States as biological control agents for the emerald ash borer (EAB), Agrilus planipennis Fairmaire, an Asian buprestid species responsible for mortality of ash trees (Fraxinus spp.) in North America. Two of these hymenopterans, Spathius agrili Yang (Braconidae), a larval ectoparasitoid, and Tetrastichus planipennisi Yang (Eulophidae), a larval endoparasitoid, prefer late-instar EAB larvae. This overlapping host preference raises concerns that interspecific competition following field releases may compromise establishment of one or both species. In a series of laboratory and field experiments, we found S. agrili and T. planipennisi exhibited similar parasitism rates when presented alone with EAB larvae for 12–14 days. However, S. agrili was more efficient at locating and parasitizing hosts within the first 27 h, possibly explaining why S. agrili excluded T. planipennisi in the laboratory trials and nearly excluded T. planipennisi in field trials when the two species were presented together with EAB larvae. We found that S. agrili parasitized larvae previously parasitized by T. planipennisi but not the reverse. However, S. agrili offspring failed to complete development on hosts that were previously parasitized by T. planipennisi. We recommend releasing these species separately in time or space to avoid the antagonistic interactions observed in this study.     

Host preference of the facultative hyperparasitoidTetrastichus howardi (Hym.: Eulophidae)

Tetrastichus howardi (Olliff) preferred the lepidopteran hosts,Chilo partellus (Swinhoe) (Pyralidae) andHelicoverpa armigera (Hübner) (Noctuidae) to their parasitoids,Xanthopimpla stemmator (Thunberg) (Hymenoptera: Ichneumonidae) andPalexorista laxa (Curran) (Diptera: Tachinidae). IfT. howardi had previously experienced parasitising a certain host, its preference for that host increased, but not significantly. When reared on a certain host, the preference for that host increased.T. howardi showed no preference to any particular age of its hosts.T. howardi was able to discriminate between parasitised and unparasitised hosts, initially preferring parasitised hosts, but two days later preferring unparasitised hosts.     

Role of emerald ash borer (Coleoptera: Buprestidae) larval vibrations in host-quality assessment by Tetrastichus planipennisi (Hymenoptera: Eulophidae)

The biological control agent Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive cambium-feeding species responsible for recent, widespread mortality of ash (Fraxinus spp.) in North America. T. planipennisi is known to prefer late-instar emerald ash borer, but the cues used to assess host size by this species and most other parasitoids of concealed hosts remain unknown. We sought to test whether vibrations produced by feeding emerald ash borer vary with larval size and whether there are any correlations between these cues and T. planipennisi progeny number (i.e., brood size) and sex ratio. The amplitudes and rates of 3-30-ms vibrational impulses produced by emerald ash borer larvae of various sizes were measured in the laboratory before presenting the larvae to T. planipennisi. Impulse-rate did not vary with emerald ash borer size, but vibration amplitude was significantly higher for large larvae than for small larvae. T. planipennisi produced a significantly higher proportion of female offspring from large hosts than small hosts and was shown in previous work to produce more offspring overall from large hosts. There were no significant correlations, however, between the T. planipennisi progeny data and the emerald ash borer sound data. Because vibration amplitude varied significantly with host size, however, we are unable to entirely reject the hypothesis that T. planipennisi and possibly other parasitoids of concealed hosts use vibrational cues to assess host quality, particularly given the low explanatory potential of other external cues. Internal chemical cues also may be important.     

Ecology,Wolbachia infection frequency and mode of reproduction in the parasitoid wasp Tetrastichus coeruleus (Hymenoptera: Eulophidae)

Whereas sexual reproduction may facilitate adaptation to complex environments with many biotic interactions, simplified environments are expected to favour asexual reproduction. In agreement with this, recent studies on invertebrates have shown a prevalence of asexual species in agricultural (simplified) but not in natural (complex) environments. We investigated whether the same correlation between reproductive mode and habitat can be found in different populations within one species. The parasitoid wasp Tetrastichus coeruleus forms an ideal model to test this question, since it occurs both in natural and agricultural environments. Further, we investigated whether Wolbachia infection caused parthenogenesis in female‐biased populations. In contrast to the general pattern, in Dutch and French natural areas, we found Wolbachia‐infected, highly female‐biased populations that reproduce parthenogenetically. In contrast, populations on Dutch agricultural fields were not infected with Wolbachia, showed higher frequencies of males and reproduced sexually. However, we also found a female‐only, Wolbachia‐infected population on agricultural fields in north‐eastern United States. All Wolbachia‐infected populations were infected with the same Wolbachia strain. At this moment, we do not have a convincing explanation for this deviation from the general pattern of ecology and reproductive mode. It may be that asparagus agricultural fields differ from other crop fields in ways that favour sexual reproduction. Alternatively, Wolbachia may manipulate life history traits in its host, resulting in different fitness pay‐offs in different habitats. The fixation of Wolbachia in the United States populations (where the species was introduced) may be due to founder effect and lack of uninfected, sexual source populations.     

Critical rearing parameters of Tetrastichus planipennisi (Hymenoptera: Eulophidae) as affected by host plant substrate and host-parasitoid group structure

In laboratory assays, we evaluated the potential impact of host plant substrate types, host-parasitoid group sizes (densities), and parasitoid-to-host ratios on select fitness parameters of the larval endoparasitoid Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae), newly introduced for biological control of emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in the United States. Results from our study showed that offspring production and critical fitness parameters (body size and sex ratio) of T. planipennisi from parasitized emerald ash borer larvae are significantly influenced by host plant substrate type, host-parasitoid group size, parasitoid-to-host ratio, or a combination in the primary exposure assay. The number of both female and male T. planipennisi progeny was significantly greater when emerald ash borer larvae were inserted into tropical ash [Fraxinus uhdei (Wenz.) Lingelsh.] logs rather than green ash (Fraxinus pensylvanica Marshall). When maintained at a constant 1:1 parasitoid-to-host ratio, assays with larger host-parasitoid group sizes (3:3-12:12) produced significantly greater numbers of both male and female offspring per parental wasp compared with those with the single host-parasitoid (1:1) group treatment. As the parasitoid-to-host ratio increased from 1:1 to 8:1 in the assay, the average brood size (number of offspring per parasitized emerald ash borer larva) increased significantly, whereas the average brood sex ratio (female to male) changed from being female-biased (6:1) to male-biased (1:2); body size of female offspring as measured by the length of ovipositor and left hind tibia also was reduced significantly. Based on these findings, we suggest that the current method of rearing T. planipennisi with artificially infested-emerald ash borer larvae use the tropical ash logs for emerald ash borer insertion, a larger (> or = 3:3) host-parasitoid group size and 1:1 parasitoid-to-host ratio in the primary parasitoid exposure assays.     

Interactions over time between cereal leaf beetle (Coleoptera: Chrysomelidae) and larval parasitoid Tetrastichus julis (Hymenoptera: Eulophidae) in Utah

The phenology of parasitism of the cereal leaf beetle, Oulema melanopus (L.) (Coleoptera: Chrysomelidae) by Tetrastichus julis (Walker) (Hymenoptera: Eulophidae) was studied in small grain fields from 2000 to 2005 in northern Utah, after release and redistribution of the partially bivoltine larval parasitoid during the 1990s. Host larvae first occurred in May, with peak infestation typically occurring in early to mid-June. Parasitism by overwintering females of T. julis was highest among earliest developing beetle larvae. Thereafter, rates of parasitism fell to low levels (5-10% or less) by the latter half of June, when heat accumulation had reached 280-350 degree-days (based on a minimum threshold of 8.9 degrees C). With the emergence of second generation parasitoids, rates of parasitism rose to levels approaching 100% among the relatively few late-developing larvae of O. inelanopus. Clear and consistent differences over the years were not observed among different crops of small grains (barley, wheat, or oats) either in the phenology and intensity of beetle infestation, or in the rate of parasitism of beetle larvae. The rate of parasitism was especially high in 2005, and an increase in the minimum level of parasitism (observed each year at mid-season) was apparent over the course of the study. These results indicate that the parasitoid has become well established and seems to be continuing to increase in its impact on O. melanopus in northern Utah, despite a relatively hostile environment of crop management, wherein most fields are plowed and disked annually.     

冈崎姬小蜂生物学特性研究

通过对福州城郊田间周年调查,发现在美洲斑潜蝇（Ｌｉｒｉｏｍｙｚａ ｓａｔｉｖａｅ Ｂｌａｎｃｈａｒｄ）寄生性天敌中,冈崎姬小蜂（Ｎｅｏｃｈｒｙｓｏｃｈａｒｉｓ ｏｋａｚａｋｉｉ Ｋａｍｉｊｏ）是主要优势种。本对冈崎姬小蜂的生物学特性进行了研究,包括交配、取食、寄生、羽化、趋性、性比、越冬、不同营养条件及不同光照时数对成蜂寿命的影响,以及各虫态历期。     

中国宽脉啮小蜂属(膜翅目:姬小蜂科)的种类及鉴定

本研究对我国姬小蜂科Eulophidae宽脉啮小蜂属Dzhanokmenia Kostjukov,1977进行分类和比较形态学研究,记录该属7种,其中中国新记录2种,提供了7种的鉴别特征及生物学和地理分布信息。比较形态学研究结果表明:宽脉啮小蜂属在头部、触角、腹部背板、腿节以及雌性外生殖器着生位置等结构上存在明显的形态特化和种间差异,这些结构特征能有效应用于属内物种鉴定,为该类群物种的准确鉴定提供依据。基于上述研究结果,本研究编制了该属的分种检索表。     

Record of Diglyphus walker (Hymenoptera: Eulophidae) species in Brazil

Leafminers (Diptera: Agromyzidae) are pests of various crops, mainly in greenhouses, and have Diglyphus spp. as important leafminer larval parasitoids. Until recently, only Diglyphus insularis (Gahan) had been reported in Brazil. In here we report the first records of Diglyphus begini (Ashmead), D. intermedius (Girault) and D. isaea (Walker) in Brazil. These parasitoids were found parasitizing leafminer larvae on cultivated and spontaneous plants in some areas of Minas Gerais state, Brazil.     

Evolution of the courtship display of Melittobia (Hymenoptera: Eulophidae)

Melittobia, a polyphagous pupal parasitoid of many solitary bees and wasps, has a complex courtship display, the evolutionary history of which is poorly understood. The current phylogeny of this genus suggests the complexity of the courtship has increased over time. We test this hypothesis using Bayesian and Maximum likelihood techniques to construct a molecular phylogeny of Melittobia-the first for this genus using modern techniques--using two intergenic regions (Internal Transcribed Spacer regions 1 and 2) and Cytochrome Oxidase 1 (CO1), and reconstruct the ancestral state of each major node in the phylogeny for five behaviors coincidental to major shifts in display complexity. We show that species-groups identified solely on behavioral characters are reliable, although the relationships among groups are different than those previously hypothesized. We also show that behaviors traditionally used in the courtship studies do not support a hypothesis of increasing display complexity over time.     

The British species of Achrysocharoides (Hymenoptera, Eulophidae)

Abstract. Taxonomic notes and a key are provided for the eleven species of Achrysocharoides found in Britain including A.carpini sp.n. Several forms of undetermined specific status are also described. Two species-groups are erected: the latreillii group and the aty s group. All British species have been reared as parasites only of leaf-mining larval Gracillariidae (Lepidoptera); one brood is here recorded from Caloptilia on Quercus; all other rearings are from Phyllono-rycter species. Parasite—host associations and speciation within the genus are discussed.