Biological control and the species status of two host-associated populations of Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae).

Abstract Populations of the morphologically defined taxon Trissolcus basalis (Wollaston) from two major hosts (Nezara viridula [L.] and Agonoscelis rutila [F.]) were tested, using biological criteria, to determine whether they represented populations of one genetical species, or whether they represented populations of two host-associated sibling species. Individuals from both hosts exhibited no obvious courtship differences and they selected mates at random in mate-choice experiments. In addition, female wasps produced offspring of both sexes after mating with a male from either population. We conclude that the individuals of the two populations tested constitute a single genetical species.     

Multiple host use by the egg parasitoid Trissolcus basalis (Wollaston) in a soybean agricultural system: biological control and environmental implications

1 Species of bugs (Hemiptera: Pentatomidae and other families) that are parasitized by the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) in the soybean agricultural system on the Darling Downs in south-eastern Queensland, Australia, are reported. The degree to which eggs of each bug species are used by T. basalis is quantified, which allows assessment of the role of these alternative host species in the biological control of the green vegetable bug Nezara viridula (L.). 2 Egg masses of nine species of pentatomid bugs and one unidentified bug species were collected. Parasitism rates of egg masses of all species averaged 50–70% but were significantly lower for the more important pest species, including the green vegetable bug, than for some of the agriculturally less important species. 3 Trissolcus basalis emerged from nearly all species collected and was the major parasitoid to emerge from most species. A number of native species were parasitized heavily by T. basalis and parasitism of such species may enhance biological control of the green vegetable bug, but may also pose environmental concerns. Parasitism of predatory bugs by T. basalis may have a negative impact on the biological control of other pests, especially lepidopterous pests. 4 Although the impact of T. basalis on native and predatory host species was not directly quantified, these host species remain abundant and therefore do not appear to be affected adversely by such high rates of parasitism.     

In vitro rearing ofTrissolcus basalis [Hym., Scelionidae] an egg parasitoid ofNezara viridula [Hem., Pentatomidae]

In vitro rearing of the egg parasitoidTrissolcus basalis (WOLL.) from eggs collected on the natural hostNezara viridula (L.) was initiated. Several oligidic diets containing insect material (Manduca sexta hemolymph or host egg content) were tested. Our initial medium with 50% hemolymph induced a high egg mortality, but by decreasing the hemolymph concentration, increasing the hen egg yolk concentration and adding 15% of free amino acids mixture, a hatching rate of 85% of the parasitoid eggs was obtained with 39% reaching the second instar and 33% the third instar. In a medium without hemolymph, but with 18% liquid from parasitized host eggs we obtained 90% to 100% hatching, 25 to 27% reaching the second instar and 8% the third instar. We did not obtain pupation from eggsin vitro, but did get pupae and adults from larvae rearedin vivo to second instar and transfered to anin vitro system.      

Influence of host species and age on host preference of Trissolcus semistriatus

Trissolcus semistriatus(Hymenoptera: Scelionidae) has a high potentialfor being an effective biological control agentfor the sunn pest, Eurygaster integriceps.This study was conducted to determine thepreferences of T. semistriatus in termsof host species and age of variousheteropteran host eggs (E. integriceps,Dolycoris baccarum, Graphosoma lineatum,Eurydema ornatum, Holcostethus vernalis). Theresults indicated that a small portion ofparasitism of E. ornatum eggs wasobtained (28.5%), although high parasitismrates were found in other host eggs(90.0–80.0%). A high percentage of adultemergence was also recorded. When tested for its ability toparasitize host eggs of different ages,T. semistriatus was found to preferyounger hosts: it showed parasitism ratesgreater than 50% with up to 3 daysold E. integriceps and D.baccarum, and with up to 4 days old G. lineatum and H. vernalis. However, theparasitism rates in all the ages of E. ornatum eggs were much lower than for other hostspecies. The developmental times in all hostspecies at different ages was extended withincreased host age. It was concluded that G. lineatum and D. baccarum could beused for mass production of egg parasitoids.     

Mating behaviour of the ‘cosmopolitan’ species Phyllognathopus viguieri (Copepoda: Harpacticoida) and its systematical significance

The mating behaviour was studied and recorded on video with individuals of four cultures of Phyllognathopus viguieri from different populations obtained from the interstitial water of a slow sand filter near the river Ruhr (Germany) (Ruhr population), from a compost heap in Bethesda (Maryland, USA) (Maryland population), from a rain gauge in Windsor Campbell farm (Jamaica) (Jamaica population), and a tree trunk with moss in a forest in the municipality of Rio de Janeiro (Brazil) (Brazil population). The mating behaviour was divided into the well‐known initial phase, copula phase and postcopulatory mate guarding phase. An additional phase prior to the initial phase serves to recognize the female, the recognition phase. The mating behaviour is identical in the males of the Jamaica and Brazil populations of P. viguieri. A postcopulatory mate guarding phase is not found in these two groups. Here, we refute the hypothesis, that a postcopulatory mate guarding phase is found in taxa in which only adult males grasp adult females. The males of the Ruhr and Maryland populations differ from each other in their mating behaviour. Generally, the males of all four populations do not mate with fertilized females which are equally unattractive to the males, i.e., females mate only once in their lifetime to produce offspring. These results corroborate the view that the different populations of P. viguieri do not belong to a single cosmopolitan species.     

Nezara viridula parasitism by the tachinid fly Trichopoda pennipes ten years after its accidental introduction into Italy from the New World

Percent parasitism of Nezara viridula (L.) (Heteroptera:Pentatomidae) by Trichopoda pennipes F.(Diptera: Tachinidae) ten years after itsfortuitous introduction was evaluated in someareas of Italy. The parasitoid is acclimatisedin central Italy with a parasitism in 1998 of21.3% in the west coast area (Lazio) and of24.8% in the inland area (Umbria). Host adultsof both sexes were attacked with the same ratein 1998, whereas this was higher in males thanfemales in 1991 and 1992. Parasitism of nymphswas lower in all three years. In 1998parasitized bugs showed 1 (54.1%) to a maximumof 15 (0.4%) macrotype eggs per individual,distributed preferentially on the thorax ascompared with other body parts. None of theother pentatomid species collected in the sameareas were parasitized by T. pennipes,indicating that there have not been indirecteffects of this fortuitous introduction tillnow.     

Seasonal variations in the encapsulation of eggs of the encyrtid parasitoid Metaphycus stanleyi by the pyriform scale, Protopulvinaria pyriformis

Encapsulation of eggs of the introduced parasitoid Metaphycus stanleyi Compere (Hymenoptera: Encyrtidae) by the pyriform scale, Protopulvinaria pyriformis (Cockerell) (Homoptera: Coccidae) under both greenhouse and field conditions, at Bet Dagan, was found to occur almost all year round (1986–1988). However, encapsulation rates varied considerably during the different seasons and were correlated with the ambient temperatures. The rates of efficient encapsulation (i) in scales infesting Hedera helix and Schefflera arboricola under greenhouse conditions, were lowest during December to May (6–17%) and highest during July to September (78–100%); (ii) in scales infesting avocado in the orchard, were lowest during October to May (0–11%) and highest during June to August (54–57%). Under greenhouse conditions, encapsulation rates did not differ in scales grown on H. helix and on S. arboricola, but were significantly lower in scales grown on avocado. Encapsulation by scales infesting S. arboricola was more frequent at temperatures ranging from 20–28 °C, than from 7–23 °C.The increased resistance by encapsulation of P. pyriformis to successful parasitization by M. stanleyi during the summer, may account for the inability of the parasitoid to prevent the autumn and winter outbreaks of the pest.     

Development and survival of Telenomus lobatus, a parasitoid of chrysopid eggs: effect of host species

Telenomus lobatus Johnson & Bin (Hymenoptera: Scelionidae), previously known only from the eggs of a berothid species, emerged from field-collected eggs of the chrysopid Mallada macleodi. In the laboratory, host species differentially influenced development, survival, and size of the parasitoid. Females parasitized between 55 and 65% of the available eggs of Chrysoperla (C. carnea, C. harrisii, and C. rufilabris), and these yielded 98 to 100% adult parasitoids within an average of 13 days. In contrast, eggs of Chrysopa were parasitized less frequently, (C. chi: 50%; C. oculata: 16%; C. quadripunctata: 1.3%) and relatively few (C. chi: 5.6%; C. oculata: 21%) or no (C. quadripunctata) parasitoids survived to adult emergence. Preimaginal development of T. lobatus in eggs of C. chi and C. oculata was approximately two days longer than in eggs of Chrysoperla spp. Although eggs of Meleoma dolicharthra were parasitized infrequently (approx. 16%), they yielded 100% adult parasitoids. Anomalochrysa maclachlani eggs were parasitized at low rates (1.8%) and parasitoid survival was intermediate (40.7%). Among all species, size of the host egg was positively related to the size of the emerging parasitoid adult.

---

Résumé T. lobatus Johnson & Bin (Hym. Scelionidae) connu jusqu'ici uniquement comme parasitoïde des ufs de Berothidae, a étè obtenu d'ufs du chrysope, Mallada macleodi récoltés dans la nature. Au laboratoire, l'espèce de l'hôte influe sur le développement, la survie et la taille du parasitoïde. Les femellles ont parasité 55 à 65% des ufs disponibles de Chrysoperla (C. carnea, C. harrisii, C. rufilabris) et ceux-ci ont donné 98 à 100% de parasitoïdes adultes en 13 jours en moyenne. Par contre, les ufs de Chrysopa étaient moins souvent parasités,-C. chi: 50%; C. oculata: 16%; C. quadripunctata: 1,3%-, et relativement peu de parasitoïdes avaient survecu jusqu'à l'émergence,-C. chi: 5,6%; C. oculata: 21%; C. quadripunctata: 0. Le développment préimaginal est approximativement 2 jours plus long dans les ufs de C. chi et C. oculata que dans ceux de Chrysoperla spp. Bien que les ufs de Meleoma dolicharthra aient été peu parasités,-environ 16%-, 100% ont donné des parasitoïdes adultes. Les ufs de Anomalochrysa maclachlani ont été peu parasités,-1,8%-, et le taux de survie a été moyen: 40,7%. Pour toutes les espèces, la taille de l'uf de l'hôte était liée positivement à la taille du parasitoïde obtenu.

     

Fecundity and Longevity of Green Vegetable Bug,Nezara viridula, Following Parasitism byTrichopoda giacomellii

The effects of parasitism by the ArgentinianTrichopoda giacomellii(Blanchard) on reproduction and longevity of its host,Nezara viridula(L.) are reported. Parasitoid larvae suppress egg maturation, reducing by 70% the fecundity of mature female hosts during the period of larval development. Egg viability was not affected, but mating frequency was reduced by approximately 50%. When parasitized as newly eclosed adults, 84% of females fail to reproduce. In male hosts, fertility and mating frequency were not affected during the period of larval parasitoid development. In male and reproductively immature female hosts, death was coincident with, or occurred shortly after parasitoid emergence (2–4 days); in mature females, death occurred on average 2 weeks after larval parasitoid emergence. Host mortality occurred as a consequence of tissue damage incurred as the parasitoid larvae emerged from the host. Some individuals survived parasitism though no further reproductive activity (mating or oviposition) occurred. The effectiveness ofT. giacomelliias a biological control agent is discussed in relation to its impact on reproduction and survival of its host and contrasted with the action of otherTrichopodaspecies.     

Identification of compounds from the eggs of Ostrinia nubilalis and Mamestra brassicae having kairomone activity on Trichogramma brassicae

In the presence of an extract of Ostrinia nubilalis or Mamestra brassicae eggs, female Trichogramma brassicae exhibited increased rates of upwind locomotion in the tubes of a linear olfactometer. GC and GC-MS analyses of O. nubilalis and M. brassicae egg extracts revealed the presence of fatty acids, their ethyl esters, and various hydrocarbons. Exposing the wasps to a mixture of the five main saturated hydrocarbons (heneicosane, tricosane, pentacosane, heptacosane and nonacosane) increased the upwind progression in the olfactometer. Single hydrocarbons elicited reduced or no activity. Ethyl palmitate and palmitic acid were also effective in increasing upwind locomotion. Z11-14:Ac, the main component of the female sex pheromone of O. nubilalis was inactive, regardless of its concentration. It is concluded that various compounds present on the surface of the O. nubilalis egg masses may play a role in the orientation of T. brassicae to its host.     

Evolutionary lag versus bill-size constraints: a comparative study of the acceptance of cowbird eggs by old hosts

Summary Evolutionary lag and resistance to puncture-ejection are two hypotheses explaining why hosts long exposed to brood parasitism by molothrine cowbirds accept their eggs. The former hypothesis assumes no physical constraints while the latter requires them. We show (1) that most acceptors of Brown-headed Cowbird eggs have small bills, (2) that the single species known to puncture cowbird eggs for ejection also has a small bill, and (3) that all grasp ejectors of Brown-headed Cowbird eggs have large bills. We propose that most acceptors cannot remove cowbird eggs by grasp ejection because their bills are too small. Small hosts probably cannot puncture cowbird eggs for ejection because their unusually thick shells make the costs of puncturing them higher than the benefits of ejection. This hypothesis is capable of explaining the success of cowbirds with scores of host species in the puzzling absence of egg mimicry.     

Flight initiation in the egg parasitoid Trichogramma minutum: Effects of ambient temperature, mates, food, and host eggs

Emergence, preening, and flight initiation were studied in laboratory-reared Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae). Male parasitoids emerged first and flew before females. When both sexes were present in flight cylinders, female parasitoids flew before males. Flight propensity in males was negatively related to the number of emerging females, while flight propensity in females was independent of the number of males present. Ambient temperature significantly affected the propensity and timing of flight; between 70–80% of the parasitoids flew at 25 and 30 °C while less than 4%, mostly males, flew at 20 °C. No flights were observed at 15 °C. The presence of fresh host eggs caused a reduction in the proportion of female parasitoids that flew and a delay in the time to flight for the females that did fly. The presence of food increased the flight propensity of female parasitoids, but did not affect the timing of flight. The relationship between flight behaviour and the efficiency of mass-reared Trichogramma is discussed in terms of its importance for inundative release programmes.     

Morphological analysis of some cryptic species in the Acanthocyclops vernalis species complex from North America

Patterns of morphological variation and reproductive isolation were examined for several North American populations of copepods in the Acanthocyclops vernalis Fischer A., 1853 (Copepoda, Cyclopinae) species complex. The copepods were collected from six sites in Wisconsin, U.S.A. Morphological analysis of 120 adult females revealed that a character used previously to distinguish species in this group was unreliable because of phenotypic plasticity. Most of the morphological variance was due to environment (Laboratory vs. field) and to field site. Relatively little of the variation was due to measurement error or asymmetry. Multivariate ordination analysis produced poorly-defined clusters of individuals, suggesting that different biological species are difficult or impossible to distinguish using a set of easily-measurable morphological characters. In our study, morphological similarity was independent of geographic distance among sites, between 0.05 and 300 km. Isofemale lines within sites showed little or no reproductive isolation, but nearly complete isolation among sites. Reproductive isolation was also independent of morphology. These results suggest that the Acanthocyclops population at each site could be considered a distinct cryptic biological species. These copepods expressed morphological stasis – persistence of morphological uniformity despite reproductive isolation. Because of the effect of site and environment on morphology, we recommend using much larger collections (many sites), common garden experiments, and a multi-disciplinary approach (morphological, reproductive, chromosomal, and molecular) as the basis for future taxonomic research on putative copepod species.     

The role of host semiochemicals in parasitoid specificity: a case study with Trissolcus brochymenae and Trissolcus simoni on pentatomid bugs

The role of semiochemicals on host specificity of two egg parasitoid species, the European Trissolcus simoni and the American Trissolcus brochymenae (Hymenoptera: Scelionidae), was studied in an olfactometer and in different arenas. Cues from two allopatric pests of cabbage, the European Eurydema ventrale and the American Murgantia histrionica (Heteroptera: Pentatomidae), and from the polyphagous and cosmopolitan Nezara viridula were tested. Both T. simoni and T. brochymenae responded to volatile and contact cues from their co-evolved hosts E. ventrale and M. histrionica, respectively, thus confirming the role of host semiochemicals in host location and recognition. When cues of non-co-evolved hosts were presented, a partial “new association” was obtained, as T. simoni probed and oviposited in M. histrionica eggs and some adult emergence occurred. However, this association is unlikely to occur in the field because T. simoni did not respond to volatile cues of M. histrionica. Instead T. brochymenae partially responded to volatile and contact cues from E. ventrale, but eggs were rarely accepted and parasitoids did not develop in this host. When N. viridula was tested, T. simoni responded only to contact cues, whereas T. brochymenae partially responded to volatile and contact cues, but N. viridula eggs were not suitable for development. Therefore, the N. viridula–T. brochymenae association reported from the literature appears unreliable. Understanding the mechanisms that result in host specificity may help increase parasitoid safety and predict their efficacy in biological control with old or new associations.     

The discovery of a new species of Ceranisus from southwestern Turkey

An explorative trip was made to southwesternTurkey in April 1995 in search of pear thrips,Taeniothrips inconsequens (Uzel)(Thysanoptera: Thripidae), and its naturalenemies. Forty-three females and 13 males of apreviously undescribed species of the genusCeranisus Walker (Hymenoptera:Eulophidae) were found, mainly in floweringArbutus andrachne L. Although theparasitoid was found in association withseveral thrips species, but especially T.inconsequens adults and larvae, its host orhosts were not determined. The parasitoid isdescribed and given the name Ceranisusantalyacus S. Triapitsyn.     

Phenology and parasitism rates in introduced populations of <Emphasis Type="Italic">Pseudacteon tricuspis</Emphasis>, a parasitoid of <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

We documented patterns of seasonal abundance and rates of parasitism in introduced populations of Pseudacteon tricuspis Borgmeier, a phorid parasitoid of the red imported fire ant, Solenopsis invicta Buren. Adult P. tricuspis populations were censused at monthly intervals for 1 year at three sites in northern Florida. Censuses were conducted by aspirating phorids attracted to disturbed S. invicta mounds. Pseudacteon tricuspis adults were present in every month at all sites, although abundances varied greatly among sites and over time. The highest densities of flies (up to 453 censused at 10 disturbed S. invicta mounds in 30 min) were observed in November, and changes in abundance over time were positively correlated among sites. Sex ratios were usually male biased. Parasitism rates were evaluated by collecting workers from field colonies and monitoring them in the laboratory for evidence of parasitism. Parasitism rates were very low – always less than 1%. The average parasitism rate per colony over 16 colonies and 2 years was 0.058%. No pupariation occurred within the first 8 days of collection, suggesting parasitism by P. tricuspis induced behavioral changes in parasitized workers that precluded such workers from our collections. If so, true field parasitism rates may be several times higher than measured here, yet still low in an absolute sense. These low parasitism rates can be reconciled with observed adult phorid densities by considering the large number of host ants present at the study sites.     

Three species of Lagenidiales isolated from the eggs and zoeae of the marine crabPortunus pelagicus

Three species of Lagenidiales were isolated from the eggs and zoeae of the marine crabPortunus pelagicus. One of them,Lagenidium callinectes, is reported for the first time in Japan, with fungal infection in the eggs.Haliphthoros milfordensis was discovered from a zoea of the crab.Atkinsiella okinawaensis was also isolated from a zoea of the crab and described as a new species. Growth temperature range and optimum temperature of the fungi were examined. All of the isolates grew at various concentrations of NaCl or KCl, and optimum growth was observed on PYGS agar containing seawater. Pathogenicity to zoeae of the swimming crabPortunus trituberculatus was demonstrated by using a zoospore suspension of each fungus at 25°C.     

Assessment ofTrichogramma species for biological control of forest lepidopteran defoliators

In a laboratory study, we determined the potential of threeTrichogramma (Hymenoptera: Trichogrammatidae) species,T. brassicae Bezdenko,T. minutum Riley andT. nr.sibiricum Sorokina, for biological control against six species of forest lepidopteran pests, black army cutworm, hemlock looper, eastern spruce budworm, western spruce budworm, white-marked tussock moth, and gypsy moth. Females of each parasitoid species were offered eggs from each of the six host species. Parasitization and the effect of the host species on the emerging progeny were examined and recorded.Trichogramma minutum had the broadest host range and successfully parasitized four host species out of the six offered.Trichogramma nr.sibiricum had the narrowest host range and parasitized only two species of hosts. Of the six host species, black army cutworm was the most preferred by all threeTrichogramma species; white-marked tussock moth and gypsy moth were not parasitized by any parasitoids. There was a positive correlation between the size of female offspring and their corresponding egg complement in all three parasitoid species. The developmental time of parasitoids from egg to adult was influenced by both the parasitoid and host species. Our results suggest thatT. minutum has the greatest potential for biological control against various forest lepidopteran pests and that the black army cutworm may be the best target candidate for further study.     

Fertility life table parameters of different strains of Trichogramma spp. collected from eggs of the carob moth Ectomyelois ceratoniae

Fertility life table parameters were assessed for five populations (strains) of two Trichogramma species, T. embryophagum Hartig and T. principium Sugonyaev & Sorokina, collected from eggs of the carob moth Ectomyelois ceratoniae (Zeller) in Iranian pomegranate orchards. Four combinations of two constant temperatures (25°C, 30°C) and two relative humidity levels (50% RH, 70% RH) were used. The overall intrinsic rate of natural increase (r_m) was significantly decreased with decreased relative humidity but was uninfluenced by temperature. The highest and lowest r_m values were observed in T. embryophagum from Qum (0.34 ± 0.004) and T. embryophagum from Varamin (0.13 ± 0.01), respectively. Two‐way interaction analyses revealed that the strains had significantly different responses in their r_m values at different temperatures and relative humidities. In general, r_m values were significantly higher at 25°C and 70% RH than at 30°C and 50% RH. The Qum strain of T. embryophagum was the most promising candidate to be considered as a biocontrol agent against E. ceratoniae due to its high reproductive rate (0.27 ± 0.01) at conditions (30°C and 50% RH) similar to the actual climatic conditions in Iranian pomegranate orchards during the major part of the growing season.     

Encapsulation of eggs of the encyrtid wasp, Metaphycus swirskii, by the hemispherical scale, Saissetia coffeae: Effects of host age and rearing temperature

The combined effects of host age and rearing temperature upon the encapsulation of eggs of Metaphycus swirskii Annecke & Mynhardt (Hymenoptera: Encyrtidae) by the hemispherical scale, Saissetia coffeae (Walker) (Homoptera: Coccidae), were determined. Encapsulation incidence at 16, 20, 24 and 28°C increased with the age of the host, being 5.5–19.6% in 3rd instar nymphs, 45.4–74.2% in young female scales and 88.8–100% in preovipositing female scales. Within each host physiological age range, encapsulation frequency was not materially influenced by most of the temperatures. At 32°C, regardless of host age, S. coffeae encapsulated 99.5 to 100% of parasitoid eggs. Complete immunity to successful parasitism by M. swirskii occurred at 20, 24, 28 and 32°C in preovipositing female scales and at 32°C in 3rd-instar nymphs and young female scales of S. coffeae.

---

Résumé Cette étude a déterminé les effets combinés de l'âge de l'hôte et de la température sur l'encapsulation des oeufs de M. swirskii Annecke & Mynhardt (Hym. Encyrtidae) par S. coffeae Walker (Homopt. Coccidae). La fréquence d'encapsulation à 16, 20, 24 et 28°C a augmenté avec l'âge de l'hôte, 5,5–19,6% dans les larves de 3ème stade, 45,4–74,2% chez les jeunes femelles et 88,8–100% chez les femelles avant la ponte. Dans chaque gamme d'âge physiologique de l'hôte, la fréquence d'encapsulation n'est pas influencée par la plupart des températures. A 32°C, quel que soit l'âge de l'hôte, S. coffeae a encapsulé 99,5–100% des oeufs du parasitoïde. Une immunité complète au parasitisme par M. swirskii a été obtenue à 20, 24, 28 et 32°C pour les femelles de cochenilles dans la période précédant la ponte, et à 32°C pour les larves de 3ème stade et les jeunes femelles de S. coffeae.