Artificial diets for rearing the coconut hispine beetle Brontispa longissima and its parasitoid Asecodes hispinarum

We tested artificial diets for rearing the coconut hispine beetle, Brontispa longissima, a serious invasive pest of coconut (Cocos nucifera) in Southeast Asia. We examined three artificial diets that were identical except for their agar content. The survival rate from hatching to adult emergence was 26.0% when beetles were reared on a ‘soft diet’ (20 g/l agar), 16.0% on a ‘hard diet’ (40 g/l agar), and 41.0% on a ‘mixed diet’, in which the hard diet was used for the first instar and the soft diet for later instars. Females raised as larvae on the mixed diet and then as adults on the soft diet laid few eggs, which did not hatch. However, females reared on the mixed diet as larvae and then on fresh leaves as adults consistently laid eggs that hatched. We then examined the suitability of B. longissima larvae reared on the mixed diet as hosts for the larval parasitoid Asecodes hispinarum, a specialist parasitoid of this beetle. Of the oviposited hosts, 75.0% became mummified and 41.7% produced adult wasps. These results indicate that the mixed diet may be useful for rearing B. longissima larvae as hosts for the rearing of A. hispinarum.     

入侵害虫椰心叶甲的研究进展

近30年来,椰心叶甲Brontispa longissima(Gestro)在亚太椰子产区扩散蔓延,成为棕榈植物上主要害虫。2002年,该害虫入侵我国海南并暴发成灾。化学防除是控制椰心叶甲危害的首要措施,其中包括挂药包等农药缓释技术,而生物防治是控制该虫扩散蔓延的经济有效措施。从感病椰心叶甲虫体上分离的微生物杀虫剂绿僵菌Metarhizium anisopliae被我国和其他多个国家应用。椰心叶甲天敌寄生蜂的引进利用是可持续控制该害虫发生的重要措施。幼虫寄生蜂椰甲截脉姬小蜂Asecodes hispinarum和蛹寄生蜂椰心叶甲啮小蜂Tetrastichus brontispae是椰心叶甲的两种重要天敌。2004年,椰甲截脉姬小蜂和椰心叶甲啮小蜂被引入到我国,经扩繁释放后,取得了良好的控害效果,其中有些地区的椰甲截脉姬小蜂寄生率可达90%,椰心叶甲啮小蜂寄生率可达100%。连续放蜂,且椰心叶甲种群有合适虫龄被寄生蜂寄生是控害效果的关键。气候等生态因子也可能是影响寄生蜂控制靶标害虫效果的因素,应进一步加强研究。论文最后提出了未来椰心叶甲防控的研究的方向。     

Artificial diets for rearing the coconut hispine beetle,Brontispa longissima (Coleoptera: Chrysomelidae), and their suitability to two specialist parasitoids

The coconut hispine beetle, Brontispa longissima (Gestro), is a serious invasive pest that infests young unopened fronds of coconut palms (Cocos nucifera L.) in Southeast Asia. We previously developed the first artificial diet for rearing B. longissima larvae, which contained a leaf powder of young coconut fronds. Because the fronds are required for healthy growth of coconut palms, it is necessary to reduce their use for rearing the beetles. In this study, we tested two new artificial diets for the beetle larvae, which contained the leaf powders of mature coconut leaves or orchard grass (Dactylis glomerata L.). Brontispa longissima successfully developed from hatching to adulthood on both the mature coconut leaf diet and orchard grass diet. The beetles reared on the mature coconut leaf diet and orchard grass diet developed faster than those reared on the young coconut leaf diet. Fecundity and egg hatchability of beetles did not differ among the three diet treatments. We then examined the suitability of beetle larvae or pupae reared on each diet as hosts for two specialist endoparasitoids, Asecodes hispinarum Boucek and Tetrastichus brontispae Ferriere. The survival rate from oviposition to adult emergence for A. hispinarum was 43.8% in hosts reared on a young coconut leaf diet, 77.1% on a mature coconut leaf diet, and 85.7% on an orchard grass diet. For T. brontispae, the survival rate was 70.0% in hosts reared on the young coconut leaf diet, 38.1% on the mature coconut leaf diet, and 66.7% on the orchard grass diet. Our results indicate these artificial diets can be useful for rearing B. longissima and its two parasitoids, helping to reduce the costs of mass rearing these insects.     

Comparative Development and Performance of Artificially Reared versus Host-RearedDiapetimorpha introita(Cresson) (Hymenoptera: Ichneumonidae) Wasps

Diapetimorpha introita(Cresson) (Hymenoptera: Ichneumonidae), a native ectoparasitoid ofSpodopteraspp. pupae, was reared in the laboratory on an artificial diet devoid of any insect host components. Diet-reared wasps demonstrated a propensity to search for and parasitize natural hosts in a field cage trial. Longevity of the diet-reared wasps was comparable with the longevity of wasps reared on host pupae. Survival rate ofD. introitawas 61.3% when reared on diet and 76.3% when reared on host pupae. Wasps reared on the artificial diet had longer developmental times, reduced fecundity, and reduced adult weights compared to wasps reared on host pupae. These studies suggest that future research efforts should focus on increasing fecundity and weight of diet-reared wasps and decreasing the mortality and developmental time of wasps reared on the artificial diet. The ability to rearD. introitaon an inexpensive, artificial diet significantly enhances the potential of mass rearing this parasitoid for inundative releases againstSpodopteraspp.     

Reproductive traits of the egg parasitoid Aprostocetus fukutai,a promising biological control agent for invasive citrus longhorned beetle Anoplophora chinensis

BioControl - Aprostocetus fukutai (Miwa & Sonan) (Hymenoptera: Eulophidae) is the only known egg parasitoid of the citrus longhorned beetle Anoplophora chinensis Forster (Coleoptera:...     

Survey of the native insect natural enemies of Hyphantria cunea (Drury) (Lepidoptera: Arctiidae) in China

The fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae), is an invasive and important pest in China. Investigations on insect natural enemies have been conducted from 1996 to 1999 in five provinces and one municipality of China in order to select effective species for biological control. Two carabid predators (Coleoptera) and 25 parasitoid species were found, among which 23 were parasitic wasps (Hymenoptera), including five hyperparasitic species and two tachinid flies (Diptera). The two carabids preyed on young larvae inside webs, two braconid wasps parasitized larvae, and 18 parasitoid species attacked the fall webworm during the pupal and/or 'larval-pupal' stages. Among these parasitoids, there were one genus and nine species that are new to science and four species new to China, which were described and published by the senior author Yang. The average parasitism rates of fall webworm pupae were 25.8% and 16.1% in the overwintering generation and the first generation (summer generation), respectively. These findings reveal that these natural enemies play an important role in the natural control of the pest. Chouioia cunea Yang (Hymenoptera: Eulophidae), a gregarious pupal endo-parasitoid, was recommended as a promising biological control agent against the fall webworm in China.     

Tritrophic interactions among coumarin, the herbivore Spodoptera litura and a gregarious ectoparasitoid Bracon hebetor

The secondary metabolites that play a defensive role in plants not only affect the growth and development of herbivores, but they can also influence their natural enemies. In the present study, the effect of coumarin was evaluated on a braconid parasitoid, Bracon hebetor (Say) (Hymenoptera: Braconidae) using polyphagous pest, Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae). Reproductive potential of the parasitoid declined significantly due to parasitization of host larvae feeding on coumarin supplemented diet. Total development period of B. hebetor extended significantly (2.17–2.38 days) when the host larvae were reared on diet amended with higher concentrations (625–3,125 parts per million) of coumarin as compared to unamended diet. Ingestion of coumarin supplemented diet by S. litura adversely affected the emergence and parasitization potential of B. hebetor. As compared to the control a significant decline in hatching of eggs was recorded when parasitoid wasps developed on host larvae fed on diet containing higher concentrations of coumarin (625–3,125 ppm). Negative effects of coumarin were also recorded from the next generation of the parasitoid.     

Phylogenetic relationships of egg parasitoids (Hymenoptera: Eulophidae) and correlated life history characteristics of their Neotropical Cassidinae hosts (Coleoptera, Chrysomelidae)

Egg parasitoids in the family Eulophidae (Hymenoptera) are an important part of the community of insects attacking neotropical leaf beetles in the subfamily Cassidinae. We present a phylogeny of 24 species of oophagous Eulophidae, using the 28S rDNA, the ITS2 rDNA and the cytochrome b genes, applying the NJ, MP, ML and Bayesian tree reconstruction methods on each data set. We ask whether the phylogenetic relationships of the parasitoids are linked with the life history characteristics of their beetle hosts. We show that cladogenesis in the oophagous Eulophidae does correlate with ovipositional behaviour and, to a lesser extent, diet and tribal affinities of their hosts. Additionally using two methods of simultaneous analysis of several gene sets: the Total Evidence method, and the construction of a "supertree" by Matrix Representation Parsimony (MRP), we substantiate the same major phylogenetic relationships within the Eulophidae.     

Using hosts of mixed sizes to reduce male-biased sex ratio in the parasitoid wasp, Diglyphus isaea

Overproduction of males in mass rearing of parasitic Hymenoptera contributes to higher costs for biological control because only females directly kill pests. We present a technique, based on manipulating host composition, to generate less male‐biased sex ratios in parasitoid species that adjust their sex allocation in response to relative host size. Our system consisted of chrysanthemum, Dendranthema grandiflora Tzvelev var. ‘Miramar’; a leafminer, Liriomyza langei Frick (Diptera: Agromyzidae); and a commercially available parasitoid, Diglyphus isaea (Walker) (Hymenoptera: Eulophidae). We compared the offspring sex ratios of D. isaea females presented with different compositions of L. langei larvae on chrysanthemum. Presenting individual females with only large hosts increased mean sex ratio from 32 to 67% male over 2 days. However, presenting individual females with progressively larger hosts over 1 or 2 days reduced mean sex ratio from 90 to 100% male to less than 30% male. Groups of females produced sex ratios around 58% male if presented with both plants infested by only small hosts and plants infested by only large hosts. In comparison, groups of females produced sex ratios around 48% male if presented with plants infested by both small hosts and large hosts. We compared the use of both small hosts and large hosts to only large hosts for simulated mass rearing of wasps over 8 weeks. Using both small hosts and large hosts produced similar numbers of wasps as using only large hosts, but reduced mean sex ratio of weekly cohorts from 66% male to 56% male. The two techniques produced females of similar size, but using both large hosts and small hosts produced slightly smaller males than using only large hosts. The use of both small hosts and large hosts for mass rearing of D. isaea could reduce actual costs of females by 23%.     

Host Location of Hyssopus pallidus, a Larval Parasitoid of the Codling Moth, Cydia pomonella

The effectiveness of parasitoids as biological control agents depends largely on their host location behavior. In this study we describe the host-searching behavior of Hyssopus pallidus (Askew), a larval parasitoid of the codling moth Cydia pomonella L. We observed parasitoid behavior on mature apples (a potential host patch) and elucidated some of the stimuli which determine host location. Female wasps showed more complex and intensive searching on infested apples than on mechanically damaged or noninfested apples. Wasps were able to enter infested apples through the calyx or the tunnel made by the host larvae and parasitize them. Area-restricted searching was observed on infested apples, in particular on areas contaminated with host frass. In a further bioassay, we confirmed that host frass contains a host location kairomone. The kairomone appears to be produced by the host independently from its diet, even though frass produced by hosts fed on an apple are more attractive than frass produced by hosts fed on a fruit-devoid artificial diet. The capability of H. pallidus to locate its host inside apples makes the foraging strategy of this species potentially useful as a biological control agent.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

Mass rearing and augmentative releases of the native parasitoid Chouioia cunea for biological control of the introduced fall webworm Hyphantria cunea in China

The fall webworm, Hyphantria cunea Drury, is an economically important introduced pest in China. A native pupal endoparasitoid, Chouioia cunea Yang (Hymenoptera: Eulophidae, Tetrastichinae) causes considerable mortality of H. cunea pupae in some areas of China. Alternative hosts of C. cunea were tested to find potential substitute hosts for mass rearing the parasitoid. Antheraea pernyi Guerin-Meneville (Lepidoptera: Saturniidae) was chosen as a surrogate host as a single pupa produced an average of 6552 wasps and a maximum of 11,256 per pupa. A mass-rearing technique and successful release program of this parasitoid is described. Parasitism by C. cunea in the areas where it was released averaged 67.74% and was usually over 80%. Other native parasitoids, such as Coccygomimus disparis (Viereck), C. parnasae (Viereck) (Hymenoptera: Ichneumonidae) and Exorita japonica Townes (Diptera: Tachnidae), caused the total average parasitism to exceed 90% on average, reaching a maximum of 96.28% in the release areas. These successful releases indicate that an introduced pest species can also be controlled by the mass-rearing and release of native parasitoids in the country of introduction.     

三种农药亚致死剂量对椰甲截脉姬小蜂功能反应的影响

测定了氯氰菊酯、啶虫脒和椰甲清3种杀虫剂亚致死剂量对椰甲截脉姬小蜂Asecodes hispinarumBoucˇek寄生椰心叶甲Brontispa logissima(Gestro)功能反应的影响。结果表明:亚致死剂量杀虫剂没有改变椰甲截脉姬小蜂功能反应模型的基本结构,但影响到了模型的各项参数。药剂处理后椰甲截脉姬小蜂的最大日捕食量降低,处理猎物时间延长,搜寻效应减弱。氯氰菊酯和啶虫脒的处理后瞬时攻击率减小,但椰甲清处理后瞬时攻击率高于对照。可见,在亚致死剂量下,氯氰菊酯和啶虫脒对椰甲截脉姬小蜂的寄生作用存在不良影响,椰甲清(30%杀虫单+15%啶虫脒)可提高其瞬时攻击率但减弱其搜索效应。     

Parasitoid dispersal and colonization lag in disturbed habitats: biological control of cereal leaf beetle metapopulations

Natural enemies of insect pests of annual crops have been hypothesized either to lag, or alternatively not to lag, behind their prey in dispersing to and colonizing new habitat. We examined parasitoid dispersal and parasitism of the cereal leaf beetle (Oulema melanopus [L.]; Coleoptera: Chrysomelidae) by the host‐specific wasp Tetrastichus julis [Walker] (Hymenoptera: Eulophidae) in wheat fields of northern Utah to assess whether a colonization lag occurred. Equally high rates of parasitism of beetle larvae (including second instars early in the year) occurred in 2010 and 2011 in fields that were newly planted to wheat vs. in fields where wheat had been grown also the previous year. A caging experiment demonstrated that parasitism in these newly planted wheat fields did not arise from parasitoid adults that had matured within the fields; instead, upon emerging in other fields, parasitoid females dispersed a minimum of 100–250 m to parasitize beetle larvae early in the spring in the newly planted fields. A transect study in 2012 revealed that T. julis females dispersed rapidly at least 600 m into a newly planted wheat field to parasitize most of the early maturing beetle larvae, which occurred at very low density. Thus, the parasitoid has very strong ability to match its host in dispersal over long distances across a highly disturbed agricultural landscape, and colonization lag appears of little importance in affecting biological control associated with this host–parasitoid interaction.     

Effects of Five Naturally Occurring Sugars on the Longevity,Oogenesis, and Nutrient Accumulation Pattern in Adult Females of the Synovigenic Parasitoid Neochrysocharis formosa (Hymenoptera: Eulophidae)

Neochrysocharis formosa (Westwood) (Hymenoptera: Eulophidae), one of the dominant natural enemies of agromyzid leafminers, is a synovigenic parasitoid. We compared the longevity, oogenesis, and nutrient levels of female wasps provided with 10% solutions of five naturally occurring sugars. All five sugars significantly increased the longevity of female wasps, which was 6.5–9.3-fold higher than that of parasitoids provided with water only. We found no significant difference in longevity of female wasps fed on glucose versus fructose or trehalose versus melezitose, but longevity of wasps fed on glucose or fructose was significantly longer than those fed on trehalose or melezitose. Also, we examined the oosorption capability of wasps fed on the five sugars. Some mature eggs were present in the ovaries of newly emerged females, but these were fully reabsorbed within 72 h when wasps were starved. Once wasps were fed with any of the sugars, the number of mature eggs increased at first and then decreased due to oosorption. The longevity and oogenesis dynamics of female wasps fed on five sugars were related with their function of hydrolysis and digestion. As female wasps have no lipogenesis capability, by acquiring exogenous sugars for oogenesis, they can either maintain or exceed the original level of capital nutrients held on adult emergence because none of the wasps’ glycogen need be metabolized to burn as sugar.     

Compound effects of induced plant responses on insect herbivores and parasitoids: implications for tritrophic interactions

1. Induced plant responses can affect herbivores either directly, by reducing herbivore development, or indirectly, by affecting the performance of natural enemies. Both the direct and indirect impacts of induction on herbivore and parasitoid success were evaluated in a common experimental system, using clonal poplar trees Populus nigra (Salicales: Salicaceae), the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae), and the gregarious parasitoid Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae). 2. Female parasitoids were attracted to leaf odours from both damaged and undamaged trees, however herbivore‐damaged leaves were three times more attractive to wasps than undamaged leaves. Parasitoids were also attracted to herbivore larvae reared on foliage and to larval frass, but they were not attracted to larvae reared on artificial diet. 3. Prior gypsy moth feeding elicited a systemic plant response that retarded the growth rate, feeding, and survival of gypsy moth larvae, however induction also reduced the developmental success of the parasitoid. 4. The mean number of parasitoid progeny emerging from hosts fed foliage from induced trees was 40% less than from uninduced trees. In addition, the proportion of parasitised larvae that survived long enough to issue any parasitoids was lower on foliage from induced trees. 5. A conceptual and analytical model is provided to describe the net impacts of induced plant responses on parasitoids, and implications for tritrophic interactions and biological control of insect pests are discussed.     

How two different host species influence the performance of a gregarious parasitoid: host size is not equal to host quality

1. Hyssopus pallidus Askew (Hymenoptera, Eulophidae) is a gregarious ectoparasitoid of the two tortricid moths species Cydia molesta Busck and C. pomonella L. (Lepidoptera, Tortricidae). It paralyses and parasitizes different larval instars of both species inside the apple fruit, which leads to the death of the caterpillar. 2. We assessed the influence of host species characteristics and host food on the performance of the parasitoid female in terms of clutch size decisions and fitness of the F(1) generation. 3. A comparison of clutch size revealed that female parasitoids deposited similar numbers of eggs on the comparatively smaller C. molesta hosts as on the larger C. pomonella hosts. The number of parasitoid offspring produced per weight unit of host larva was significantly higher in C. molesta than in C. pomonella, which is contrary to the general prediction that smaller hosts yield less parasitoid offspring. However, the sex ratio was not influenced by host species that differed considerably in size. 4. Despite the fact that less host resources were available per parasitoid larva feeding on C. molesta caterpillars, the mean weight of emerging female wasps was higher in the parasitoids reared on C. molesta. Furthermore, longevity of these female wasps was neither influenced by host species nor by the food their host had consumed. In addition we did not find a positive relationship between adult female weight and longevity. 5. Parasitoid females proved to be able to assess accurately the nutritional quality of an encountered host and adjust clutch size accordingly. These findings indicate that host size is not equal to host quality. Thus host size is not the only parameter to explain the nutritional quality of a given host and to predict fitness gain in the subsequent generation.     

Classical biological control of the cereal leaf beetle,Oulema melanopus (Coleoptera: Chrysomelidae), in Washington State and rôle of field insectaries,a review

The cereal leaf beetle (CLB) Oulema melanopus (Coleoptera: Chrysomelidae), was considered a high risk to Washington State’s cereal grain production when the pest was found there in 1999. Biological control agents, which had proven successful in the Midwest, were introduced beginning in 2000. The parasitoids were released into field insectaries that were modified for the region and set up at CLB hotspots across the state. The egg parasitoid, Anaphes flavipes (Förster) (Hymenoptera: Mymaridae), failed to establish. The larval parasitoid, Tetrastichus julis (Walker) (Hymenoptera: Eulophidae), established successfully at all insectaries. Assays of commercial grain fields showed the parasitoid dispersing widely beyond the insectaries. A proactive Extension programme enabled farmers to benefit from the biocontrol without applying insecticides to manage the CLB. A modified insectary, consisting of oat strips seeded between commercial fields of winter and spring wheat, demonstrated potential as a way for farmers to increase parasitoid populations on their land. Within 5 years of first overwintering, T. julis had established across the state and was effectively suppressing CLB.     

Superparasitism and host size effects in Oomyzus sokolowskii, a parasitoid of diamondback moth

Many aspects of a parasitoid's biology may be affected by its host. Host size, for example, could affect parasitoid fitness, especially in gregarious parasitoids, in which the resource is used by multiple siblings. Oomyzus sokolowskii (Kurdjumov) (Hymenoptera: Eulophidae) is a gregarious larval–pupal endoparasitoid of the diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), a major pest of crucifers worldwide, and is able to superparasitize the host. This study focuses on the hypothesis that because resource availability is higher in larger hosts, parasitoids developing in larger hosts will fare better. However, superparasitized hosts are expected to yield larger numbers of parasitoid offspring of smaller body size. Results showed that superparasitism increased the number of parasitoid offspring produced per host and increased offspring longevity, but decreased offspring body size. However, developmental time and sex ratio of parasitoid offspring was similar among hosts parasitized once, twice, or three times. Regardless of superparasitism, parasitoids emerging from larger hosts that were fed honey solution lived longer than similarly fed progeny from smaller hosts (36.4 vs. 22.1 days). The results partially support the hypothesis that Oomyzus gained fitness from an increase in host size; moreover, superparasitism seems advantageous for Oomyzus due to increased offspring numbers and longevity.     

Factors affecting the parasitism rate and the number and sex ratio of offspring of Asecodes hispinarum Bouček (Hymenoptera: Eulophidae), a biological control agent of Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae)

To optimise the production of Asecodes hispinarum Bou?ek (Hymenoptera: Eulophidae), a parasitoid of coconut leaf beetle, Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae), some of the factors affecting rates of parasitism, number of offspring produced per host and sex ratio of A. hispinarum were investigated. The numbers and sex ratio of A. hispinarum offspring per host reduced significantly at extreme low humidity (30% relative humidity [RH]), but there was no significant effect on parasitism. Photoperiod had no significant effects on any of the life traits tested. A. hispinarum was able to reproduce via arrhenotoky, and while increasing the proportion of female parents increased the number of parasitoids produced, the proportion of female offspring decreased. Older females showed a lower rate of parasitism than young females, however, maternal age did not affect the number or the sex ratio of offspring. Increasing the number of hosts offered to a pair of parasitoids significantly increased the number of parasitised hosts but decreased the parasitism rate while the sex ratio of progeny was not affected. Present work showed that to maximise the production of female parasitoids, a parasitoid/host ratio of 1:1, using one-day old A. hispinarum at a female/male ratio of 3:1 and RH of at least 55% is recommended.