Sex allocation theory reveals a hidden cost of neonicotinoid exposure in a parasitoid wasp.

Sex allocation theory has proved to be one the most successful theories in evolutionary ecology. However, its role in more applied aspects of ecology has been limited. Here we show how sex allocation theory helps uncover an otherwise hidden cost of neonicotinoid exposure in the parasitoid wasp Nasonia vitripennis. Female N. vitripennis allocate the sex of their offspring in line with Local Mate Competition (LMC) theory. Neonicotinoids are an economically important class of insecticides, but their deployment remains controversial, with evidence linking them to the decline of beneficial species. We demonstrate for the first time to our knowledge, that neonicotinoids disrupt the crucial reproductive behaviour of facultative sex allocation at sub-lethal, field-relevant doses in N. vitripennis. The quantitative predictions we can make from LMC theory show that females exposed to neonicotinoids are less able to allocate sex optimally and that this failure imposes a significant fitness cost. Our work highlights that understanding the ecological consequences of neonicotinoid deployment requires not just measures of mortality or even fecundity reduction among non-target species, but also measures that capture broader fitness costs, in this case offspring sex allocation. Our work also highlights new avenues for exploring how females obtain information when allocating sex under LMC.

Pheromone diversification and age-dependent behavioural plasticity decrease interspecific mating costs in Nasonia.

Interspecific mating can cause severe fitness costs due to the fact that hybrids are often non-viable or less fit. Thus, theory predicts the selection of traits that lessen reproductive interactions between closely related sympatric species. Males of the parasitic wasp Nasonia vitripennis differ from all other Nasonia species by an additional sex pheromone component, but the ecological selective forces underlying this pheromone diversification are unknown. Here we present data from lab experiments suggesting that costly interspecific sexual interactions with the sympatric species N. giraulti might have been responsible for the pheromone evolution and some courtship-related behavioural adaptations in N. vitripennis. Most N. giraulti females are inseminated already within the host, but N. giraulti males still invest in costly sex pheromones after emergence. Furthermore, they do not discriminate between N. vitripennis females and conspecifics during courtship. Therefore, N. vitripennis females, most of which emerge as virgins, face the risk of mating with N. giraulti resulting in costly all-male broods due to Wolbachia-induced cytoplasmic incompatibility. As a counter adaptation, young N. vitripennis females discriminate against N. giraulti males using the more complex conspecific sex pheromone and reject most of them during courtship. With increasing age, however, N. vitripennis females become less choosy, but often compensate mating errors by re-mating with a conspecific. By doing so, they can principally avoid suboptimal offspring sex ratios, but a microcosm experiment suggests that under more natural conditions N. vitripennis females cannot completely avoid fitness costs due to heterospecific mating. Our study provides support for the hypothesis that communication interference of closely related sympatric species using similar sexual signals can generate selective pressures that lead to their divergence.

Effects of spirotetramat on Aonidiella aurantii (Homoptera: Diaspididae) and its parasitoid, Aphytis melinus (Hymenoptera: Aphelinidae).

Laboratory and field studies were conducted to measure the effects of spirotetramat on life stages of California red scale, Aonidiella aurantii (Maskell), and a primary parasitoid, Aphytis melinus DeBach. Organophosphate-resistant and -susceptible populations responded similarly to spirotetramat, suggesting there is no cross-resistance between these insecticide classes. First and second instar male and female A. aurantii were 10- and 32-fold more susceptible to spirotetramat (LC50 = 0.1-0.2 ppm) compared with early third (LC50 = 1.5 ppm) and late third instar females (LC50 = 5.3 ppm). The LC99 value indicated that late stage third instar females would not be fully controlled by a field rate of spirotetramat; however, spirotetramat would reduce their fecundity by 89%. Field applications of spirotetramat in two water volumes and using two adjuvants (oil and a nonionic spray adjuvant) showed similar reduction in A. aurantii numbers, even though the higher water volume demonstrated more complete coverage. These data suggest that this foliarly applied systemic insecticide can be applied in as little as 2,340 liters/ha of water volume, minimizing application costs, and that the two adjuvants acted similarly. The endoparasitoid, A. melinus, was unaffected by the field rate of spirotetramat when it was applied to the host when the parasitoid was in the egg or larval stage. Adult A. melinus showed 2 wk of moderate reductions in survival when exposed to leaves with field-weathered residues. Spirotetramat is an integrated pest management compatible insecticide, effective in reducing A. aurantii stages and allowing survival of its primary parasitoid A. melinus.

Risk assessment of selected insecticides on Tamarixia triozae (Hymenoptera: Eulophidae), a parasitoid of Bactericera cockerelli (Hemiptera: Trizoidae).

Tamarixia triozae (Burks) (Hymenoptera: Eulophidae) is an important parasitoid of the potato or tomato psyllid, Bactericera cockerelli (Sulc) (Hemiptera: Trizoidae), a serious pest of potato (Solanum tuberosum L.), tomato (Solanum lycopersicum L.), and other solanaceous vegetables in many countries. To produce a marketable crop, insecticides are required when B. cockerelli populations reach economically damaging levels. We evaluated 11 commonly used insecticides for their effects on T. triozae. Glass-surface residues of spinetoram, imidacloprid-cyfluthrin, abamectin, and tolfenpyrad caused 100% mortality of T. triozae in 72 h, and the leaf residue of spinetoram was extremely toxic to T. triozae adults; even 15-d-old residues caused 100% mortality. Cyantraniliprole, fenpyroximate, pymetrozine, spirotetramat, spiromesifen, and chenopodium oil did not cause significant mortality in either glass surface or leaf-residue bioassays. Ingestion of spinetoram, abamectin, and imidacloprid+cyfluthrin (Leverage) by the adults resulted in 100% mortality in 12 h, and tolfenpyrad, 75.0% mortality in 12 h; whereas chenopodium oil and pymetrozine showed moderate effects on adult survival. Ingestion of abamectin, imidacloprid-cyfluthrin, and spinetoram killed all adults in the first day of treatment, whereas female adults in the treatment of pymetrozine lived 80.8 d, which was similar to those in the control. Ingestion of abamectin, imidacloprid-cyfluthrin, chenopodium oil, and spinetoram killed all male adults in the first day, whereas ingestion of other insecticides did not cause significant mortality, but reduced percent parasitism. Abamectin, imidacloprid-cyfluthrin, and spinetoram had the most deleterious effects on T. triozae, and have the least potential for use in integrated control programs using this parasitoid.

Assessment of toxicity risk of insecticides used in rice ecosystem on Trichogramma japonicum, an egg parasitoid of rice lepidopterans.

Both chemical and biological methods are essential for control of insects, for example, lepidopterans, on rice. Thus, it is important to know the effect of chemicals on the biological control agents. In this study, we assessed the toxicity of commonly used insecticides on a biological control agent, Trichogramma japonicum Ahmead (an egg parasitoid of rice lepidopterans) by using a dry film residue method. Results showed that thirty insecticides from seven chemical classes exhibited various degree of toxicity to this parasitoid. Among the seven classes of chemicals tested, organophosphates (chlorpyrifos, fenitrothion, phoxim, profenofos, and triazophos) and carbamates (carbaryl, carbsulfan, isoprocarb, metolcarb, and promecarb) exhibited the highest intrinsic toxicity to T. japponicum, with an LC50 of 0.035 (0.029-0.044) to 0.49 (0.34-0.87) mg active ingredient (a.i.) L(-1), followed by antibiotics (abamectin, emamectin benzoate, and ivermectin), phenylpyrazoles (butane-fipronil, ethiprole, and fipronil), pyrethroids (cyhalthrin, cypermethrin, fenpropathrin, and lambda-cyhaothrin), and neonicotinoids (acetamiprid, imidacloprid, imidaclothiz, nitenpyram, thiacloprid, and thiamethoxam). Moreover, the insect growth regulator insecticides (chlorfluazuron, fufenozide, hexaflumuron and tebufenozide) exhibited the lowest toxicity to the wasps with an LC50 of 3,383 (2406-5499) to 30206 (23107-41008) mg ai. L(-1). Risk quotient analysis showed that phenylpyrazoles, pyrethroids, insect growth regulators, neonicotinoids (with the exception of thiamethoxam), and antibiotics (with the exception of abamectin) are classified as safe agents to the parasitoid, while organophosphates and carbamates are classified as slightly, moderately, or highly toxic agents to the parasitoid. The data presented in this paper provided useful information on the selection of compatible insecticides with T. japonicum.

Insecticides suppress natural enemies and increase pest damage in cabbage.

Intensive use of pesticides is common and increasing despite a growing and historically well documented awareness of the costs and hazards. The benefits from pesticides of increased yields from sufficient pest control may be outweighed by developed resistance in pests and killing of beneficial natural enemies. Other negative effects are human health problems and lower prices because of consumers' desire to buy organic products. Few studies have examined these trade-offs in the field. Here, we demonstrate that Nicaraguan cabbage (Brassica spp.) farmers may suffer economically by using insecticides as they get more damage by the main pest diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), at the same time as they spend economic resources on insecticides. Replicated similarly sized cabbage fields cultivated in a standardized manner were either treated with insecticides according common practice or not treated with insecticides over two seasons. Fields treated with insecticides suffered, compared with nontreated fields, equal or, at least in some periods of the seasons, higher diamondback moth pest attacks. These fields also had increased leaf damage on the harvested cabbage heads. Weight and size of the heads were not affected. The farmers received the same price on the local market irrespective of insecticide use. Rates of parasitized diamondback moth were consistently lower in the treated fields. Negative effects of using insecticides against diamondback moth were found for the density of parasitoids and generalist predatory wasps, and tended to affect spiders negatively. The observed increased leaf damages in insecticide-treated fields may be a combined consequence of insecticide resistance in the pest, and of lower predation and parasitization rates from naturally occurring predators that are suppressed by the insecticide applications. The results indicate biological control as a viable and economic alternative pest management strategy, something that may be particularly relevant for the production of cash crops in tropical countries where insecticide use is heavy and possibly increasing.