Cotesia Sesamiae Research Articles

Uncertainties in the effectiveness of biological control of stem borers under different climate change scenarios in Eastern Africa

Climate change (CC) is expected to significantly affect biodiversity and ecosystem services. Adverse impacts from CC in the Global South are likely to be exacerbated by limited capacities to take adequate adaptation measures and existing developmental challenges. Insect pests today are already causing considerable yield losses in agricultural crop production in East Africa. Studies have shown that insects are strongly responding to CC by proliferation, shift in distribution, and by altering their phenology, which is why an impact on agriculture can be expected. Biological control (BC) has been proposed as an alternative measure to sustainably contain insect pests, but few studies predict its efficacy under future CC. Using the species maximum entropy modeling (Maxent) approach, we predict the current and future distribution of three important lepidopteran stem borer pests of maize in Eastern Africa, i.e., Busseola fusca (Fuller, 1901), Chilo partellus (Swinhoe, 1885), and Sesamia calamistis (Hampson, 1910), and two parasitoids that are currently used for BC, i.e., Cotesia flavipes (Cameron, 1891) and Cotesia sesamiae (Cameron, 1906). Based on these potential distributions and data collected during household surveys with local farmers in Kenya and Tanzania, also future maize yield losses are predicted for a business-as-usual scenario and a sustainable development scenario. We found that BC of the stem borer pests by C. flavipes and C. sesamiae will be less effective under more severe CC resulting in a reduced ability to curb maize yield losses caused by the stem borers. These results highlight the need to adapt BC measures to future CC to maintain its potential for environmentally friendly pest management strategies. The findings of this research are thus of particular relevance to policymakers, extension officers, and farmers in the region and will aid the adaptation of smallholder agricultural practices to the impacts of CC.

Biogeography of cereal stemborers and their natural enemies: forecasting pest management efficacy under changing climate.

Climate warming presents physiological challenges to insects, manifesting as loss of key life-history fitness traits and survival. For interacting host-parasitoid species, physiological responses to heat stress may vary, thereby potentially uncoupling trophic ecological relationships. Here, we assessed heat tolerance traits and sensitivity to prevailing and future maximum temperatures for the cereal stemborer pests, Chilo partellus, Busseola fusca and Sesamia calamistis and their endo-parasitoids, Cotesia sesamiae and Cotesia flavipes. We further used the machine learning algorithm, Maximum Entropy (MaxEnt), to model current and potential distribution of these species. The mean critical thermal maxima (CTmax ) ranged from 39.5 ± 0.9°C to 44.6 ± 0.6°C and from 46.8 ± 0.7°C to 48.5 ± 0.9°C for parasitoids and stemborers, with C. sesamiae and Ch. partellus exhibiting the lowest and highest CTmax respectively. From the current climate to the 2050s scenario, parasitoids recorded a significant reduction in warming tolerance compared with their hosts. Habitat suitability for all stemborer-parasitoid species was spatially heterogeneous under current and future climatic scenarios. Cotesia sesamiae C. flavipes and B. fusca exhibited significant habitat loss, whereas Ch. partellus and S. calamistis showed a significant habitat gain under future 2050s predictions. Model metrics based on mean area under the curve ranged from 0.72 to 0.84 for all species, indicating a good predictive performance of the models. These results suggest C. sesamiae and C. flavipes may face survival constraints or extirpation compared with their pest hosts when environmental temperature reaches their upper thermal limits earlier, likely reducing pest regulation through density-mediated effects. The results demonstrate potential destabilization of stemborer-parasitoid trophic systems potentially compromising biocontrol efficacy under climate warming. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Natural Enemies of Fall Armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) in Different Agro-Ecologies.

Simple SummaryFall armyworm (FAW), Spodoptera frugiperda, was reported in Cameroon for the first time in 2017 and by the end of 2018; the pest was found all over the country. Cameroon is among the first countries in Africa where the southern armyworm (SAW), Spodoptera eridania, another economical important armyworm was reported. The African governments adopted emergency actions around chemical insecticides despite the range of economic and health risks associated with chemical control. This work aims at identifying parasitoids (natural enemies) of armyworms and test their acceptability, suitability, and host range on Spodoptera spp., that can play a significant role in the sustainable management of these spodopterans. Field surveys conducted lead to the identification of two egg and four larval parasitoids. The fall armyworm was the predominant spodopteran. Laboratory studies were conducted to assess the performance of parasitoids associated with both pests in Cameroon. T. remus showed significantly higher parasitism on FAW than SAW, with significantly shorter development time on FAW, while inducing significant non-reproductive mortality on FAW. Laboratory performance of larval parasitoid was not compared between the two spodopterans identified but the developmental parameters showed that C. icipe has a shorter development time compared to other larval parasitoids.Fall armyworm (FAW) Spodoptera frugiperda (J.E. Smith) and southern armyworm (SAW) Spodoptera eridania (Stoll) have become major threats to crops in Africa since 2016. African governments adopted emergency actions around chemical insecticides, with limited efforts to assess the richness or roles of indigenous natural enemies. Field surveys and laboratory studies were conducted to identify and assess the performance of parasitoids associated with spodopterans in Cameroon. FAW was the most abundant spodopteran pest. Telenomus remus (Nixon), Trichogramma chilonis (Ishi), Charops sp. (Szépligeti), Coccygidium luteum (Cameron), Cotesia icipe (Fernandez & Fiaboe), and Cotesia sesamiae (Cameron) are the first records in the country on spodopterans. Telenomus remus, T. chilonis, C. icipe, and Charops sp. were obtained from both FAW and SAW; C. luteum and C. sesamiae from FAW. The distribution of spodopterans, their endoparasitoids, and parasitism rates varied with host, season and location. In the laboratory, T. remus showed significantly higher parasitism on FAW than SAW, and significant differences in the development parameters between the two host eggs, with shorter development time on FAW. It induced significant non-reproductive mortality on FAW but not on SAW. Developmental parameters showed that C. icipe has a shorter development time compared to other larval parasitoids. Implications for conservative and augmentative biocontrol are discussed.

Managing and monitoring genetic isolation and local adaptation of endemic and introduced Cotesia sesamiae for the biological control of the cereal stemborer Busseola fusca in Cameroon

The success of biological bontrol (BC) introductions can be enhanced by considering theory and knowledge of biological systems. The gregarious braconid parasitoid Cotesia sesamiae (Cameroon) is one of the best studied biological control agent from the perspective of molecular ecology. Its evolutionary adaptation to the target host involves symbiotic partners. Polydnaviruses are responsible for immune and developmental adaptations whereas Wolbachia bacteria may reinforce this local adaptation though genetic isolation mechanisms. The noctuid Busseola fusca is a major stemborer pest of maize in sub-Saharan Africa. In contrast to eastern Africa, C. sesamiae is rarely found on B. fusca in western Africa. It is however often obtained from other stemborer species feeding on wild grasses. A biological control project was launched in 2006–2007 by introducing to Cameroon seven crosses of Kenyan populations of C. sesamiae collected in different ecozones. They included populations adapted to B. fusca that develop on maize as well as populations adapted to other hosts feeding on wild plants to allow carryover between cropping seasons. Wolbachia strains responsible for cytoplasmic reproductive incompatibilities with endemic strains were included in the crosses to limit genetic exchanges between introduced and endemic C. sesamiae and preserve genetic adaptation to B. fusca of the introduced populations, while at the same time preserving their ability to survive on wild plants. Six post release surveys were carried out on maize from 2007 to 2013, and on wild grasses in 2013. A total of 393 C. sesamiae individuals, each from one cocoon mass, were genotyped for 11 microsatellite loci. Multidimensional scaling analysis, STRUCTURE and GENECLASS analyses assigned almost all the parasitoids recovered from maize to those introduced from Kenya. The introduced strains were also recovered from wild host plants with little genetic exchanges with endemics. Each population remained strongly associated with its original Wolbachia component, suggesting that Wolbachia may contribute to genetic isolation between endemics and introduced populations in wild host plants when maize is absent, thereby conciliating biological control success and safety.

Impact of the exotic fall armyworm on larval parasitoids associated with the lepidopteran maize stemborers in Kenya

Exotic invasive insect herbivores have the potential to interfere with existing herbivore-natural enemy interactions in new environments. Fall armyworm (FAW), Spodoptera frugiperda J.E. Smith (Lepidoptera: Noctuidae), is a new invasive pest in maize fields in Africa. Understanding the acceptability and suitability of FAW to existing maize stemborer-parasitoid interactions is the first step in elucidating the impact that this exotic insect pest can have on the existing natural enemies used in biological control of maize stemborers in Kenya. The most commonly used larval parasitoids for biological control programs against maize stemborer communities in East Africa are Cotesia flavipes Cameron (Hymenoptera: Braconidae) and two populations of the native Cotesia sesamiae (Cs-Inland and Cs-Coast) Cameron (Hymenoptera: Braconidae). All these parasitoid species attacked FAW larvae but none yielded offspring, although they induced high non-reproductive host mortality when compared to natural mortality. Furthermore, the parasitoids that inserted their ovipositor into FAW larvae exhibited no significant preference between FAW larvae and their respective stemborer hosts under dual-choice bioassays. In olfactometer bioassays, the parasitoids were more attracted to plants infested by FAW than uninfested plants and even showed a marked preference for the odours of plants infested by FAW over those of plants infested by their natural host counterparts. This study illustrates that exotic pests, such as FAW, can impact existing stemborer-parasitoid interactions associated with maize, even if they cannot be used as hosts by parasitoids associated with these stemborers. Although additional studies are needed, FAW might therefore have a negative impact on stemborer biological control existing before its invasion.

Genome wide association analysis of a stemborer egg induced \u201ccall-for-help\u201d defence trait in maize

Tritrophic interactions allow plants to recruit natural enemies for protection against herbivory. Here we investigated genetic variability in induced responses to stemborer egg-laying in maize Zea mays (L.) (Poaceae). We conducted a genome wide association study (GWAS) of 146 maize genotypes comprising of landraces, inbred lines and commercial hybrids. Plants were phenotyped in bioassays measuring parasitic wasp Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) attraction to volatiles collected from plants exposed to stemborer Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) eggs. Genotyping-by-sequencing was used to generate maize germplasm SNP data for GWAS. The egg-induced parasitoid attraction trait was more common in landraces than in improved inbred lines and hybrids. GWAS identified 101 marker-trait associations (MTAs), some of which were adjacent to genes involved in the JA-defence pathway (opr7, aos1, 2, 3), terpene biosynthesis (fps3, tps2, 3, 4, 5, 7, 9, 10), benzoxazinone synthesis (bx7, 9) and known resistance genes (e.g. maize insect resistance 1, mir1). Intriguingly, there was also association with a transmembrane protein kinase that may function as a receptor for the egg elicitor and other genes implicated in early plant defence signalling. We report maize genomic regions associated with indirect defence and provide a valuable resource for future studies of tritrophic interactions in maize. The markers identified may facilitate selection of indirect defence by maize breeders.

Relative Influence of Host, Wolbachia, Geography and Climate on the Genetic Structure of the Sub-saharan Parasitic Wasp Cotesia sesamiae

The parasitoid lifestyle represents one of the most diversified life history strategies on earth. There are however very few studies on the variables associated with intraspecific diversity of parasitoid insects, especially regarding the relationship with spatial, biotic and abiotic ecological factors. Cotesia sesamiae is a Sub-Saharan stenophagous parasitic wasp that parasitizes several African stemborer species with variable developmental success. The different host-specialized populations are infected with different strains of Wolbachia, an endosymbiotic bacterium widespread in arthropods that is known for impacting life history traits, notably reproduction, and consequently species distribution. In this study, first we analyzed the genetic structure of C. sesamiae across Sub-Saharan Africa, using 8 microsatellite markers. We identified five major population clusters across Sub-Saharan Africa, which probably originated in the East African Rift region and expanded throughout Africa in relation to host genus and abiotic factors, such as Koppen-Geiger climate classification. Using laboratory lines, we estimated the incompatibility between the different strains of Wolbachia infecting C. sesamiae. We observed that incompatibility between Wolbachia strains was asymmetric, expressed in one direction only. Based on these results, we assessed the relationships between the direction of gene flow and Wolbachia infections in the genetic clusters. We found that host specialization was more influential on genetic structure than Wolbachia-induced reproductive incompatibility, which in turn was more influential than geography and current climatic conditions. These results are discussed in the context of African biogeography, and co-evolution between Wolbachia, virus parasitoid and host, in the perspective of improving biological control efficiency through a better knowledge of biological control agents’ evolutionary ecology.

Carry-Over Niches for Lepidopteran Maize Stemborers and Associated Parasitoids during Non-Cropping Season.

Sources of infestation are the key elements to be considered in the development of habitat management techniques for the control of maize stemborers. Several wild plants, grasses mostly, have been identified that serve as hosts for stemborers and their parasitoids during the off-season when maize is not present in the field. However, their abundance is much lower in wild plants compared to cultivated fields. Thus, the role of wild plants as a reservoir for cereal stemborers and their parasitoids is still controversial, particularly in agro-ecosystems with reduced wild habitat. We studied the occurrence of different maize stemborers and associated parasitoids in maize stem residues and wild grasses during non-cropping seasons as potential carry-over populations to subsequent early season maize plants. Surveys were conducted in the central region of Kenya during long and short dry seasons in maize residues and wild grasses as well as during the two rainy seasons in maize plants at earlier and late whorl stages during the years of 2017 and 2018. Wild habitat had a higher species diversity than maize residues habitat, but maize residues had a higher abundance of maize stemborer species, such as Busseola fusca, Sesamia calamistis, and Chilo partellus, and of associated parasitoid species (i.e., Cotesia flavipes and Cotesia sesamiae) than wild plants. Our surveys, complemented by field parasitoid releases of C. flavipes and C. sesamiae, indicated that maize residues constitute a better refugia reservoir not only of the maize stemborers but also of C. flavipes and C. sesamiae during non-cropping seasons as compared to wild plants and, thus, might constitute in this region the main source of both stemborers and C. flavipes/C. sesamiae carry-over in maize plants during the subsequent cropping season. Thus, systematic destruction of maize residues would not help the biological control of lepidopteran stemborers. This is particularly true in areas with reduced wild habitat.

Genetic footprints of adaptive divergence in the bracovirus of Cotesia sesamiae identified by targeted resequencing

The African parasitoid wasp Cotesia sesamiae is a generalist species structured in locally adapted populations showing differences in host range. The recent discovery of Cotesia typhae, a specialist, sister species to C.sesamiae, provides a good framework to study the genetic determinants of parasitoid host range. To investigate the genomic bases of divergence between these populations and species, we used a targeted sequencing approach on 24 samples. We targeted the bracovirus genomic region encoding virulence genes involved in the interaction with the lepidopteran hosts of the wasps. High sequencing coverage was obtained for all samples, allowing the study of genetic variation between wasp populations and species. By combining population genetic estimations, such as nucleotide diversity (π), relative differentiation (FST ) and absolute divergence (dxy ), with branch-site dN/dS measures, we identified six of 98 bracovirus genes showing significant divergence and evidence of positive selection. These genes, belonging to different gene families, are potentially involved in host adaptation and in the specialization process. Fine-scale analyses of genetic variation also revealed mutations and large deletions in certain genes inducing pseudogenization and loss of function. The image emerging from these results is that adaptation mediated by bracovirus genes happens through selection of particularly adaptive alleles and loss of nonadaptive genes. These results highlight the central role of the bracovirus in the molecular interactions between the wasps and their hosts and in the evolutionary processes of specialization.

Tri-Trophic Impacts of Bt-Transgenic Maize on Parasitoid Size and Fluctuating Asymmetry in Native vs. Novel Host-Parasitoid Interactions in East Africa.

Environmental stress can affect trait size and cause an increase in the fluctuating asymmetry (FA) of bilateral morphological traits in many animals. For insect parasitoids, feeding of hosts on transgenic maize, expressing a Bacillus thuringiensis toxin gene is a potential environmental stressor. We compared the size of antennae, forewings, and tibia, as well as their FA values, in two parasitoids developed on two East African host species feeding on non-transgenic vs. transgenic maize. The two lepidopteran stem-borer hosts were the native Sesamia calamistis Hampson (Lepidoptera: Noctuidae) and a recent invader, Chilo partellus Swinhoe (Lepidoptera: Crambidae). The two braconid parasitoids were the native, gregarious larval endoparasitoid Cotesia sesamiae and the recently introduced Cotesia flavipes. Both parasitoids attacked both hosts, creating evolutionarily old vs. novel interactions. Transient feeding of hosts on transgenic maize had various effects on FA, depending on trait as well as the host and parasitoid species. These effects were usually stronger in evolutionarily novel host–parasitoid associations than in the older, native ones. These parameters have capacity to more sensitively indicate the effects of potential stressors and merit further consideration.

The Cotesia sesamiae story: insight into host-range evolution in a Hymenoptera parasitoid and implication for its use in biological control programs.

This review covers nearly 20years of studies on the ecology, physiology and genetics of the Hymenoptera Cotesia sesamiae, an African parasitoid of Lepidoptera that reduces populations of common maize borers in East and South Africa. The first part of the review presents studies based on sampling of C. sesamiae from maize crops in Kenya. From this agrosystem including one host plant and three main host borer species, studies revealed two genetically differentiated populations of C. sesamiae species adapted to their local host community, and showed that their differentiation involved the joint evolution of virulence genes and sensory mechanisms of host acceptance, reinforced by reproductive incompatibility due to Wolbachia infection status and natural inbreeding. In the second part, we consider the larger ecosystem of wild Poales plant species hosting many Lepidoptera stem borer species that are potential hosts for C. sesamiae. The hypothesis of other host-adapted C. sesamiae populations was investigated based on a large sampling of stem borer larvae on various Poales across sub-Saharan Africa. The sampling provided information on the respective contribution of local hosts, biogeography and Wolbachia in the genetic structure of C. sesamiae populations. Molecular evolution analyses highlighted that several bracovirus genes were under positive selection, some of them being under different selection pressure in C. sesamiae populations adapted to different hosts. This suggests that C. sesamiae host races result from co-evolution acting at the local scale on different bracovirus genes. The third part considers the mechanisms driving specialization. C. sesamiae host races are more or less host-specialized. This character is crucial for efficient and environmentally-safe use of natural enemies for biological control of pests. One method to get an insight in the evolutionary stability of host-parasite associations is to characterize the phylogenetic relationships between the so-called host-races. Based on the construction of a phylogeny of C. sesamiae samples from various host- and plant species, we revealed three main lineages. Mechanisms of differentiation are discussed with regard to the geography and ecology of the samples. One of the lineage presented all the hallmarks of a distinct species, which has been morphologically described and is now studied in the perspective of being used as biological control agent against Sesamia nonagrioides Lefèbvre (Lepidoptera: Noctuidae), a major maize pest in West Africa and Mediterranean countries (see Benoist et al. 2017). The fourth part reviews past and present use of C. sesamiae in biological control, and points out the interest of such molecular ecology studies to reconcile biodiversity and food security stakes in future biological control.

Comparative assessment of the thermal tolerance of spotted stemborer, Chilo partellus (Lepidoptera: Crambidae) and its larval parasitoid, Cotesia sesamiae (Hymenoptera: Braconidae).

Under stressful thermal environments, insects adjust their behavior and physiology to maintain key life-history activities and improve survival. For interacting species, mutual or antagonistic, thermal stress may affect the participants in differing ways, which may then affect the outcome of the ecological relationship. In agroecosystems, this may be the fate of relationships between insect pests and their antagonistic parasitoids under acute and chronic thermal variability. Against this background, we investigated the thermal tolerance of different developmental stages of Chilo partellus Swinhoe (Lepidoptera: Crambidae) and its larval parasitoid, Cotesia sesamiae Cameron (Hymenoptera: Braconidae) using both dynamic and static protocols. When exposed for 2 h to a static temperature, lower lethal temperatures ranged from -9 to 6 °C, -14 to -2 °C, and -1 to 4 °C while upper lethal temperatures ranged from 37 to 48 °C, 41 to 49 °C, and 36 to 39 °C for C. partellus eggs, larvae, and C. sesamiae adults, respectively. Faster heating rates improved critical thermal maxima (CTmax ) in C. partellus larvae and adult C. partellus and C. sesamiae. Lower cooling rates improved critical thermal minima (CTmin ) in C. partellus and C. sesamiae adults while compromising CTmin in C. partellus larvae. The mean supercooling points (SCPs) for C. partellus larvae, pupae, and adults were -11.82 ± 1.78, -10.43 ± 1.73 and -15.75 ± 2.47, respectively. Heat knock-down time (HKDT) and chill-coma recovery time (CCRT) varied significantly between C. partellus larvae and adults. Larvae had higher HKDT than adults, while the latter recovered significantly faster following chill-coma. Current results suggest developmental stage differences in C. partellus thermal tolerance (with respect to lethal temperatures and critical thermal limits) and a compromised temperature tolerance of parasitoid C. sesamiae relative to its host, suggesting potential asynchrony between host-parasitoid population phenology and consequently biocontrol efficacy under global change. These results have broad implications to biological pest management insect-natural enemy interactions under rapidly changing thermal environments.

An Indirect Defence Trait Mediated through Egg-Induced Maize Volatiles from Neighbouring Plants.

Attack of plants by herbivorous arthropods may result in considerable changes to the plant’s chemical phenotype with respect to emission of herbivore-induced plant volatiles (HIPVs). These HIPVs have been shown to act as repellents to the attacking insects as well as attractants for the insects antagonistic to these herbivores. Plants can also respond to HIPV signals from other plants that warn them of impending attack. Recent investigations have shown that certain maize varieties are able to emit volatiles following stemborer egg deposition. These volatiles attract the herbivore’s parasitoids and directly deter further oviposition. However, it was not known whether these oviposition-induced maize (Zea mays, L.) volatiles can mediate chemical phenotypic changes in neighbouring unattacked maize plants. Therefore, this study sought to investigate the effect of oviposition-induced maize volatiles on intact neighbouring maize plants in ‘Nyamula’, a landrace known to respond to oviposition, and a standard commercial hybrid, HB515, that did not. Headspace volatile samples were collected from maize plants exposed to Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) egg deposition and unoviposited neighbouring plants as well as from control plants kept away from the volatile emitting ones. Behavioural bioassays were carried out in a four-arm olfactometer using egg (Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)) and larval (Cotesia sesamiae Cameron (Hymenoptera: Braconidae)) parasitoids. Coupled Gas Chromatography-Mass Spectrometry (GC-MS) was used for volatile analysis. For the ‘Nyamula’ landrace, GC-MS analysis revealed HIPV production not only in the oviposited plants but also in neighbouring plants not exposed to insect eggs. Higher amounts of EAG-active biogenic volatiles such as (E)-4,8-dimethyl-1,3,7-nonatriene were emitted from these plants compared to control plants. Subsequent behavioural assays with female T. bournieri and C. sesamiae parasitic wasps indicated that these parasitoids preferred volatiles from oviposited and neighbouring landrace plants compared to those from the control plants. This effect was absent in the standard commercial hybrid we tested. There was no HIPV induction and no difference in parasitoid attraction in neighbouring and control hybrid maize plants. These results show plant-plant signalling: ‘Nyamula’ maize plants emitting oviposition-induced volatiles attractive to the herbivore’s natural enemies can induce this indirect defence trait in conspecific neighbouring undamaged maize plants. Maize plants growing in a field may thus benefit from this indirect defence through airborne signalling which may enhance the fitness of the volatile-emitting plant by increasing predation pressure on herbivores.

Ongoing ecological speciation in Cotesia sesamiae, a biological control agent of cereal stem borers.

To develop efficient and safe biological control, we need to reliably identify natural enemy species, determine their host range, and understand the mechanisms that drive host range evolution. We investigated these points in Cotesia sesamiae, an African parasitic wasp of cereal stem borers. Phylogenetic analyses of 74 individual wasps, based on six mitochondrial and nuclear genes, revealed three lineages. We then investigated the ecological status (host plant and host insect ranges in the field, and host insect suitability tests) and the biological status (cross-mating tests) of the three lineages. We found that one highly supported lineage showed all the hallmarks of a cryptic species. It is associated with one host insect, Sesamia nonagrioides, and is reproductively isolated from the other two lineages by pre- and postmating barriers. The other two lineages had a more variable phylogenetic support, depending on the set of genes; they exhibited an overlapping and diversified range of host species and are not reproductively isolated from one another. We discuss the ecological conditions and mechanisms that likely generated this ongoing speciation and the relevance of this new specialist taxon in the genus Cotesia for biological control.

Chemical cues modulating electrophysiological and behavioural responses in the parasitic waspCotesia sesamiae

A better understanding of the chemical cues employed by parasitoids to locate their herbivore hosts will contribute towards effective exploitation of parasitoids as biological control agents against damaging crop pests. The current study was conducted to establish key bioactive compounds mediating behavioural response in the parasitic wasp Cotesia sesamiae (Cameron, 1906) (Hymenoptera: Braconidae) by employing electrophysiological techniques and behavioural studies. Electroantennogram (EAG) recording with synthetic equivalents of herbivore-induced plant volatiles (HIPVs) using C. sesamiae confirmed that 13 compounds elicited electrophysiological activity. In an olfactometer bioassay, only (E)-4,8-dimethyl-1,3,7-nonatriene, (E)-β-farnesene, and (E,E)-4,8,-trimethyl-1,3,7-tridecatetraene elicited behavioural responses from the parasitoid when tested individually at a natural dose, while (R)-linalool and (E)-caryophyllene were attractive at the highest test dose. Other EAG-active HIPVs were not attractive either at lower or higher concentrations, suggesting their role as blend components rather than functioning as attractants by themselves. In contrast, nine-component synthetic HIPV blends formulated in the same ratio and concentration as in a natural sample was as attractive as the original headspace sample. By providing further insights into maize–stemborer–parasitoid tritrophic interactions, results from this study will help in selecting crop cultivars emitting appropriate HIPVs to attract natural enemies and perhaps repel damaging crop pests.

Diversity of stem borer parasitoids and their associated wild host plants in South Africa and Mozambique

The diversity of lepidopterous stem borers, their parasitoids and their associated wild host plants was studied in South Africa between 2006 and 2009 and in Mozambique between 2005 and 2010. In South Africa, 20 species of parasitoids were recovered from 17 stem borer species collected on 16 wild host plant species. From Mozambique, 14 parasitoid species were recorded from 16 stem borer species collected on 14 wild host plant species. The highest diversity of parasitoids was recorded on stem borers that attacked the host plants Phragmites australis (7 spp.) and Panicum maximum (6 spp.), in South Africa and Mozambique, respectively. Bracon sp. (Hymenoptera: Braconidae) and Procerochasmias nigromaculatus (Hymenoptera: Ichneumonidae) were the most recorded parasitoid species in South Africa while Cotesia sesamiae (Hymenoptera: Braconidae) parasitized most stem borer species in Mozambique. The most common tachinid parasitoid recorded in this study was Sturmiopsis parasitica (Diptera: Tachinidae). Parasitism of stem borers during the off season was previously thought to occur mainly in natural habitats but this study shows that although natural habitats provided refuges for some parasitoid species, stem borer parasitism was generally low in wild host plants, irrespective of whether collections were done during the cropping or off-season. Parasitoid beta diversity did not depend on habitat or host plant species, but was determined by stem borer diversity.

Oviposition Induced Volatile Emissions from African Smallholder Farmers’ Maize Varieties

Maize (corn), Zea mays, is a genetically diverse crop, and we have recently shown that certain open pollinated varieties (OPVs) of Latin American origin possess a trait not present in mainstream commercial varieties: they produce volatiles in response to stemborer oviposition that are attractive to stemborer parasitoids. Here, we tested whether a similar tritrophic effect occurs in the African OPVs 'Nyamula' and 'Jowi'. Herbivore induced plant volatiles (HIPVs) were collected from plants exposed to egg deposition by the stemborer Chilo partellus. In a four-arm olfactometer bioassay, the parasitic wasp Cotesia sesamiae preferred samples containing HIPVs from plants with eggs to samples collected from plants without eggs. EAG-active compounds, including (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), were released in higher amounts from the egg induced headspace samples. Our results suggest that this oviposition trait is not limited to S. American Z. mays germplasm, and that it could be used to increase indirect defense against attack by stemborers.

Maize landraces recruit egg and larval parasitoids in response to egg deposition by a herbivore

Natural enemies respond to herbivore-induced plant volatiles (HIPVs), but an often overlooked aspect is that there may be genotypic variation in these 'indirect' plant defence traits within plant species. We found that egg deposition by stemborer moths (Chilo partellus) on maize landrace varieties caused emission of HIPVs that attract parasitic wasps. Notably, however, the oviposition-induced release of parasitoid attractants was completely absent in commercial hybrid maize varieties. In the landraces, not only were egg parasitoids (Trichogramma bournieri) attracted but also larval parasitoids (Cotesia sesamiae). This implies a sophisticated defence strategy whereby parasitoids are recruited in anticipation of egg hatching. The effect was systemic and caused by an elicitor, which could be extracted from egg materials associated with attachment to leaves. Our findings suggest that indirect plant defence traits may have become lost during crop breeding and could be valuable in new resistance breeding for sustainable agriculture.

Intraspecific specialization of the generalist parasitoid Cotesia sesamiae revealed by polyDNAvirus polymorphism and associated with different Wolbachia infection

As a result of an intense host-parasite evolutionary arms race, parasitic wasps frequently display high levels of specialization on very few host species. For instance, in braconid wasps very few generalist species have been described. However, within this family, Cotesia sesamiae is a generalist species that is widespread in sub-Saharan Africa and develops on several lepidopteran hosts. In this study, we tested the hypothesis that C.sesamiae may be a cryptic specialist when examined at the intraspecific level. We sequenced exon 2 of CrV1, a gene of the symbiotic polyDNAvirus that is integrated into the wasp genome and is associated with host immune suppression. We found that CrV1 genotype was more closely associated with the host in which the parasitoid developed than any abiotic environmental factor tested. We also tested a correlation between CrV1 genotype and an infection with Wolbachia bacteria, which are known for their ability to induce reproductive isolation. The Wolbachia bacteria infection polymorphism was also found as a major factor explaining the genetic structure of CrV1, and, in addition, the best model explaining CrV1 genetic structure involved an interaction between Wolbachia infection and host species. We suggest that Wolbachia could act as an agent capable of maintaining advantageous alleles for host specialization in different populations of C.sesamiae. This mechanism could be applicable to other insect models because of the high prevalence of Wolbachia in insects.

Sensilla on antennae, ovipositor and tarsi of the larval parasitoids, Cotesia sesamiae (Cameron 1906) and Cotesia flavipes Cameron 1891 (Hymenoptera: Braconidae): a comparative scanning electron microscopy study

Two braconid parasitoids of cereal stemborers in eastern Africa, Cotesia sesamiae and Cotesia flavipes, have been shown to display a similar hierarchy of behavioural events during host recognition and acceptance. In order to understand the mechanisms underlying host recognition and acceptance, the morphology of antennal sensilla on the last antennomeres, on the ovipositor, and on the fifth tarsomere and pretarsus of the prothoracic legs tarsi were studied using scanning electron microscopy followed by selective silver nitrate staining. It appeared that female C. sesamiae and C. flavipes shared the same types and distribution of sensory receptors, which enable them to detect volatiles and contact chemical stimuli from their hosts. In both parasitoids, four types of sensilla were identified on the three terminal antennomeres: (i) non-porous sensilla trichodea likely to be involved in mechanoreception, (ii) uniporous sensilla chaetica with porous tips that have gustatory functions, (iii) multiporous sensilla placodea, which are likely to have olfactory function, and (iv) sensilla coeloconica known to have thermo-hygroreceptive function. The tarsi of both parasitoids possessed a few uniporous sensilla chaetica with porous tips, which may have gustatory functions. The distal end of the ovipositor bore numerous dome-shaped sensilla. However, there were no sensilla coeloconica or styloconica, known to have gustatory function in other parasitoid species, on the ovipositors of the two braconid wasps.