Cotesia Species Research Articles

Wolbachia strain diversity in a complex group of sympatric cryptic parasitoid wasp species

BackgroundMaternally-inherited symbionts can induce pre-mating and/or post-mating reproductive isolation between sympatric host lineages, and speciation, by modifying host reproductive phenotypes. The large parasitoid wasp genus Cotesia (Braconidae) includes a diversity of cryptic species, each specialized in parasitizing one to few related Lepidoptera host species. Here, we characterized the infection status of an assemblage of 21 Cotesia species from 15 countries by several microbial symbionts, as a first step toward investigating whether symbionts may provide a barrier to gene flow between these parasitoid host lineages.ResultsThe symbiotic microbes Arsenophonus, Cardinium, Microsporidium and Spiroplasma were not detected in the Cotesia wasps. However, the endosymbiotic bacterium Wolbachia was present in at least eight Cotesia species, and hence we concentrated on it upon screening additional DNA extracts and SRAs from NCBI. Some of the closely related Cotesia species carry similar Wolbachia strains, but most Wolbachia strains showed patterns of horizontal transfer between phylogenetically distant host lineages.ConclusionsThe lack of co-phylogenetic signal between Wolbachia and Cotesia suggests that the symbiont and hosts have not coevolved to an extent that would drive species divergence between the Cotesia host lineages. However, as the most common facultative symbiont of Cotesia species, Wolbachia may still function as a key-player in the biology of the parasitoid wasps. Its precise role in the evolution of this complex clade of cryptic species remains to be experimentally investigated.

Two new European species of Cotesia Cameron, 1891 (Hymenoptera: Braconidae, Microgastrinae) parasitizing butterflies (Lepidoptera: Papilionoidea), and an unrelated synonymy in the genus

Two new European species of Cotesia are described and illustrated. One (C. euchloevora Shaw, sp. nov.) was reared from Euchloe species and close relatives and also Aplocera efformata, and the other (C. parnassii Shaw, sp. nov.) was reared from Parnassius phoebus. Notes are given to distinguish them from congeners. The DNA barcodes of C. euchloevora are discussed in relation to similar barcodes for another, but morphologically clearly distinct, species Cotesia pilicornis (Thomson). Unrelated to this, it is proposed that C. acutula (Tobias) is a junior synonym of C. cajae (Bouché).

Synopsis of the parasitoid wasp genus Cotesia Cameron, 1891 (Hymenoptera: Braconidae: Microgastrinae) in Australia, with the description of seven new species

The genus Cotesia Cameron, 1891 is one of the most diverse of the Microgastrinae, a subfamily of wasps that are exclusively endoparasitic on lepidopteran larvae. Species of Cotesia are widely utilised as biological control agents across the world. In Australia, there are currently 10 confirmed native species as well as four species introduced for the management of lepidopteran pests. The genus is morphologically conserved and has not been studied in the Australasian region for many decades. In this study, we use both comparative morphology and sequence data from the COI gene to delineate species, and in so doing describe seven new species from Australia: C. lasallei sp. nov., C. medusae sp. nov., C. ocellata sp. nov., C. reidarum sp. nov., C. scripta sp. nov., C. tjapekki sp. nov. and C. wonboynensis sp. nov., raising the number of species of Cotesia formally recorded in Australia to 21. We also provide updated descriptions of the previously described native species, diagnoses for the introduced species and a key to all currently described species found on the continent and from Papua New Guinea. This study treats only a fraction of the likely diversity of Cotesia, but provides a solid framework for future work.

Distinct Roles of Cuticular Aldehydes as Pheromonal Cues in Two Cotesia Parasitoids.

Cuticular compounds (CCs) that cover the surface of insects primarily serve as protection against entomopathogens, harmful substances, and desiccation. However, CCs may also have secondary signaling functions. By studying the role of CCs in intraspecific interactions, we may advance our understanding of the evolution of pheromonal communication in insects. We previously found that the gregarious parasitoid, Cotesia glomerata (L.), uses heptanal as a repellent pheromone to help avoid mate competition among sibling males, whereas another cuticular aldehyde, nonanal, is part of the female-produced attractive sex pheromone. Here, we show that the same aldehydes have different pheromonal functions in a related solitary parasitoid, Cotesia marginiventris (Cresson). Heptanal enhances the attractiveness of the female's sex pheromone, whereas nonanal does not affect a female's attractiveness. Hence, these common aldehydes are differentially used by the two Cotesia species to mediate, synergistically, the attractiveness of the main constituents of their respective sex pheromones. The specificity of the complete sex pheromone blend is apparently regulated by two specific, less volatile compounds, which evoke strong electroantennographic (EAG) responses. This is the first demonstration that volatile CCs have evolved distinct pheromonal functions to aid divergent mating strategies in closely related species. We discuss the possibility that additional compounds are involved in attraction and that, like the aldehydes, they are likely oxidative products of unsaturated cuticular hydrocarbons.

Evolutionary relationships of courtship songs in the parasitic wasp genus, Cotesia (Hymenoptera: Braconidae).

Acoustic signals play an important role in premating isolation based on sexual selection within many taxa. Many male parasitic wasps produce characteristic courtship songs used by females in mate selection. In Cotesia (Hymenoptera: Braconidae: Microgastrinae), courtship songs are generated by wing fanning with repetitive pulses in stereotypical patterns. Our objectives were to sample the diversity of courtship songs within Cotesia and to identify e underlying patterns of differentiation. We compared songs among 12 of ca. 80 Cotesia species in North America, including ten species that have not been recorded previously. For Cotesia congregata, we compared songs of wasps originating from six different host-foodplant sources, two of which are considered incipient species. Songs of emergent males from wild caterpillar hosts in five different families were recorded, and pattern, frequency, and duration of song elements analyzed. Principal component analysis converted the seven elements characterized into four uncorrelated components used in a hierarchical cluster analysis and grouped species by similarity of song structure. Species songs varied significantly in duration of repeating pulse and buzz elements and/or in fundamental frequency. Cluster analysis resolved similar species groups in agreement with the most recent molecular phylogeny for Cotesia spp., indicating the potential for using courtship songs as a predictor of genetic relatedness. Courtship song analysis may aid in identifying closely related cryptic species that overlap spatially, and provide insight into the evolution of this highly diverse and agriculturally important taxon.

Cotesia icipe sp. n., a new Microgastrinae wasp (Hymenoptera, Braconidae) of importance in the biological control of Lepidopteran pests in Africa

A new species of Microgastrinae, Cotesia icipe Fernández-Triana & Fiaboe, sp. n., is described from eastern Africa. It was reared in Kenya as a solitary parasitoid from two major amaranth pests, Spodoptera littoralis (Boisduval, 1833) and S. exigua (Hübner, 1808); study of specimens in collections also revealed its presence in four other countries in the Afrotropical region (Madagascar, Saudi Arabia, South Africa, and Yemen). Morphological, molecular and biological characters are used to describe the new species and to distinguish it from all 12 previously described species of Afrotropical Cotesia. Cotesia icipe shows potential in the biological control of key Lepidopteran pests in small scale farming conditions in Africa.

Combined use of herbivore‐induced plant volatiles and sex pheromones for mate location in braconid parasitoids

Herbivore-induced plant volatiles (HIPVs) are important cues for female parasitic wasps to find hosts. Here, we investigated the possibility that HIPVs may also serve parasitoids as cues to locate mates. To test this, the odour preferences of four braconid wasps - the gregarious parasitoid Cotesia glomerata (L.) and the solitary parasitoids Cotesia marginiventris (Cresson), Microplitis rufiventris Kokujev and Microplitis mediator (Haliday) - were studied in olfactometers. Each species showed attraction to pheromones but in somewhat different ways. Males of the two Cotesia species were attracted to virgin females, whereas females of M. rufiventris were attracted to virgin males. Male and female M. mediator exhibited attraction to both sexes. Importantly, female and male wasps of all four species were strongly attracted by HIPVs, independent of mating status. In most cases, male wasps were also attracted to intact plants. The wasps preferred the combination of HIPVs and pheromones over plant odours alone, except M. mediator, which appears to mainly use HIPVs for mate location. We discuss the ecological contexts in which the combined use of pheromones and HIPVs by parasitoids can be expected. To our knowledge, this is the first study to show that braconid parasitoids use HIPVs and pheromones in combination to locate mates.

A review of unusual species of Cotesia (Hymenoptera, Braconidae, Microgastrinae) with the first tergite narrowing at midlength.

The unusual species of Cotesia (Hymenoptera, Braconidae, Microgastrinae) with the first tergite narrowing at midlength are reviewed. One new species, Cotesia trabalae sp. n. is described from India and compared with Cotesia pistrinariae (Wilkinson) from Africa, the only other species sharing the same character of all the described species worldwide. The generic placement of these two species, based on molecular and morphological analyses as well as parasitoid biology is discussed.

Molecular Evidence on Symbiotic Relationships of Bracovirus and Cotesia Species Based on PTP R Region

Molecular Evidence on Symbiotic Relationships of Bracovirus and Cotesia Species Based on PTP R Region

Parasitoid Wasp Cotesia congregata (Say) (Insecta: Hymenoptera: Braconidae)

This species of Cotesia has been widely used as a model system in insect physiology. It has also been used to examine insect learning in host-parasitoid-plant interactions. It is an important natural enemy of the tobacco hornworm, a detrimental pest species that feeds on many plants in the Solanaceae (tobacco, pepper, tomato, etc.) family. This 5-page fact sheet was written by Christopher D. Crockett, Andrea Lucky, Oscar E. Liburd, and Karen M. Kester, and published by the UF Department of Entomology and Nematology, July 2014.
 EENY598/IN1042: A Parasitoid Wasp Cotesia congregata (Say) (Insecta: Hymenoptera: Braconidae) (ufl.edu)

Synopsis of Cotesia species, biological agents for pest control on vegetables in Vietnam, with description of one new species

The biology of four allied species of the genusCotesiaCameron is reviewed. All these species were reared from larvae of pest insects mainly belonging to the families Lymantriidae, Noctuidae, Pieridae and Plutellidae. They are larval endoparasitoids and considered as important agents for biological control of pest insects infesting different vegetable crops in Vietnam, such as cabbage, kohlrabi, small colza, cauliflower, peanut, soybeans, and various other legumes. Of those four species,Cotesia clethrogynaeLong sp. n., is described as new and illustrated; the biology and distribution of the other three allied species are given, they are illustrated and compared, and a key to four species is provided.This paper resulted from the identification of a new species ofCotesiareared from limantrid larvae ofClethrogynae turbataButler and the necessity to compare related species. The host is an important pest infestant on soybean, different beans, peas and peanut in Vietnam.

Convergence and Divergence in Direct and Indirect Life-History Traits of Closely Related Parasitoids (Braconidae: Microgastrinae)

Closely related species in nature often show similarities in suites of direct and indirect traits that reveal aspects of their phylogenetic history. Here we tested how common descent affects trait evolution in several closely related parasitoid species in the genera Cotesia and Microplitis (Hymenoptera: Braconidae: Microgastrinae) by comparing development, resource use and allocation into reproduction and maintenance. Parasitoids in these genera exhibit traits, like haemolymph feeding as larvae and external pupation that are rare in most parasitoid lineages. The growth of parasitized hosts was reduced by 90 % compared with healthy hosts, and maximum host size depended to a large extent on adult parasitoid size. Development time was longer in the more generalist parasitoids than in the specialists. Adult body mass was sexually dimorphic in all Cotesia species, with females being larger, but not in Microplitis spp. In contrast, in one of the Microplitis species males were found to be the larger sex. Egg load dynamics during the first 6 days after emergence were highly variable but egg number was typically higher in Cotesia spp. compared to Microplitis spp. Longevity in the various species was only greater in female than in male wasps in two Microplitis sp. There was a clear inverse relationship between resource use and allocation, e.g. maximum egg load and longevity, in these parasitoids. Our results reveal that adaptation to constraints imposed by host quality and availability has resulted in trait convergence and divergence at the species, genus and subfamily level.

Identification of a Cotesia species parasitizing on <I>Pieris rapae</I> from Beijing

Identification of a Cotesia species parasitizing on <I>Pieris rapae</I> from Beijing

Intrinsic competition among solitary and gregarious endoparasitoid wasps and the phenomenon of ‘resource sharing’

1. Intrinsic competition was compared in three species of braconid wasps, the solitary Meteorus pulchricornis Wesmael, and the gregarious Cotesia kariyai (Watanabe) and Cotesia ruficrus Haliday in caterpillars of their common host, the armyworm Mythimna separata Walker. Competition was determined in pair‐wise contests consisting of simultaneous and subsequent parasitisms at various time intervals between the first and second attacks (<1, 24, 48, 72, and 96 h).2. The experiment revealed that the solitary M. pulchricornis parasitoid was generally superior over the other two gregarious Cotesia species, although, in some cases C. kariyai out‐competed M. pulchricornis when ovipositing first. In contrast, irrespective of the time interval between parasitism and multiparasitism, C. ruficrus always lost in competition with M. pulchricornis.3. Remarkably, both C. kariyai and C. ruficrus occasionally emerged from the same multiparasitised caterpillar when the time interval between the first and second attacks was small. However, cumulative brood sizes of both species were generally less than when C. kariyai and C. ruficrus developed alone, suggesting that some competition did occur between these two species.4. Inter‐specific ‘resource sharing’ amongst parasitoid larvae has rarely been described in the literature. In the present study, the conditions in which ‘resource sharing’ in parasitoids can evolve are discussed. Further, we describe differing host usage strategies in parasitoids and how these can affect the outcome of aggressive larval competition.

Parasitism of Lepidopterous Stem Borers in Cultivated and Natural Habitats

Plant infestation, stem borer density, parasitism, and parasitoid abundance were assessed during two years in two host plants, Zea mays (L.) (Cyperales: Poaceae) and Sorghum bicolor (L.) (Cyperales: Poaceae), in cultivated habitats. The four major host plants (Cyperus spp., Panicum spp., Pennisetum spp., and Sorghum spp.) found in natural habitats were also assessed, and both the cultivated and natural habitat species occurred in four agroecological zones in Kenya. Across habitats, plant infestation (23.2%), stem borer density (2.2 per plant), and larval parasitism (15.0%) were highest in maize in cultivated habitats. Pupal parasitism was not higher than 4.7% in both habitats, and did not vary with locality during each season or with host plant between each season. Cotesia sesamiae (Cameron) and C. flavipes Cameron (Hymenoptera: Braconidae) were the key parasitoids in cultivated habitats (both species accounted for 76.4% of parasitized stem borers in cereal crops), but not in natural habitats (the two Cotesia species accounted for 14.5% of parasitized stem borers in wild host plants). No single parasitoid species exerted high parasitism rates on stem borer populations in wild host plants. Low stem borer densities across seasons in natural habitats indicate that cereal stem borer pests do not necessarily survive the non-cropping season feeding actively in wild host plants. Although natural habitats provided refuges for some parasitoid species, stem borer parasitism was generally low in wild host plants. Overall, because parasitoids contribute little in reducing cereal stem borer pest populations in cultivated habitats, there is need to further enhance their effectiveness in the field to regulate these pests.

Molecular evidence for the presence of the endosymbiontic bacteria Wolbachia in Cotesia populations (Hymenoptera: Braconidae)

Cotesia are larval endoparasitoids and are potential biocontrol agents for various insect pests. In the present study, we attempted to detect the allele-specific ftsZ gene of Wolbachia in Cotesia spp. obtained from different geographical locations. Samples of Cotesia plutellae originating from India (Bangalore and Ludhiana), Kenya, and South Africa and samples of C. glomerata from India (Solan) confirmed the presence of Wolbachia. However, C. plutellae samples from the Kullu region of India did not show infection with Wolbachia. ftsZ sequences showed a high degree of homology within the species (> 99%). The low genetic distance in the infected Cotesia populations indicated a single ancestral type, and distance to and from the outgroup suggested that populations are from the same species. Phylogenetic grouping of Cotesia species on the basis of geographic origin showed that the Wolbachia strain is closely related to strains that infect phylogenetically distant hosts.

Geographic distribution, host range and perennation of Cotesiasesamiae and Cotesia flavipes Cameron in cultivated and natural habitats in Kenya

Field surveys were carried out in four agroecological zones to assess the geographic distribution, host range and perennation of Cotesia sesamiae and Cotesia flavipes in cultivated and natural habitats in Kenya. The distribution of the two Cotesia species in different ecological regions was most affected by the suitability of the local stem borer species for parasitoid development, and temperature, as both species were found in localities dominated by their suitable host(s) where temperature favoured their occurrence. Fourteen years after its release, C. flavipes has maintained a high level of specificity to its target host Chilo partellus on maize and sorghum in cultivated habitats and on Sorghum arundinaceum in natural habitats. Cotesia flavipes appeared to be an appropriate biological control agent against C. partellus in eastern Africa, with minimal or no effects on non-target hosts in different habitats. Conversely, C. sesamiae lacked host specificity in different habitats, as its stem borers or host plants varied with both locality and habitat type. Perennation by both Cotesia species occurred mainly in cultivated habitats. Furthermore, natural habitats played a role in sustaining some individuals of C. flavipes during both rainy and dry seasons. These areas acted as refuges for C. flavipes, but not for C. sesamiae, because its hosts were scarce on natural host plants. The availability of these Cotesia species across seasons was mainly influenced by the presence of actively feeding stem borers on cereal plants during different seasons, as well as the duration of the dry season in different localities.

Investment in cocoon-silk and structure of the clusters of cocoons produced by gregarious microgastrine wasps (Hymenoptera: Braconidae)

After leaving their hosts, the larvae of endoparasitic braconid wasps pupate in cocoons. To determine their investment in cocoon silk, the dry weight of newly emerged wasps and that of the empty cocoons were measured for three gregarious braconid species of slightly different sizes: Glyptapanteles liparidis (Bouche), Cotesia glomerata (L.) and Cotesia kariyai (Watanabe) (Hymenoptera: Braconidae, Microgastrinae). These braconids form clusters of cocoons of different types. Glyptapanteles liparidis is significantly larger than either of the Cotesia species, and C. kariyai is the smallest. The ratio of the weight of cocoon silk to the total weight of cocoon silk, wasp body, cast cuticle and meconium is smaller for small species than large species. Small species econo- mise on their use of silk by aggregating cocoons and can therefore invest a larger fraction of their resources in adult body mass. Moreover, the larvae of the smallest species, C. kariyai, additionally reduce their use of silk by constructing a communal airy silk layer beneath which the individual cocoons are formed.

Resource allocation and bionomics of indigenous and exoticCotesia(Hymenoptera: Braconidae) species reared onSesamia calamistis

The braconid larval parasitoids Cotesia chilonis (Matsumura), C. flavipes Cameron and a strain of Cotesia sesamiae (Cameron) from coastal Kenya, reared at the International Centre of Insect Ecology and Physiology, were introduced at the International Institute of Tropical Agriculture in the Republic of Benin for suitability testing on West African stemborers prior to release. C. chilonis was originally collected in Japan while C. flavipes was imported into Kenya from Pakistan. The host species used was the noctuid Sesamia calamistis (Hampson), the most important noctuid maize pest in the region. All three Cotesia species attacked and successfully developed in 2nd to 6th larval instar of S. calamistis but parasitoid-induced mortality was highest on second instars. On most instars, C. sesamiae and C. flavipes produced larger broods than C. chilonis. Larvae parasitized by C. sesamiae developed to the 6th instar and attained an average larval weight of 353 mg, while larvae parasitized by C. chilonis only molted to the 4th instar and attained a maximum weight of 107 mg. The lower developmental threshold estimated from the non-linear regression of temperature on developmental rate was 15.9, 15.9 and 14.9 degrees C for C. chilonis, C. sesamiae and C. flavipes, respectively, while the maximum temperature was 34.2, 35.2 and 33.8 degrees C, respectively. A maximum of four ovipositions were observed per female during a life span ranging from 1.3 days for C. chilonis and C. flavipes to 1.6 days for C. sesamiae. The largest adult progeny, intrinsic rate of increase and net reproductive rates were recorded at 28 degrees C for all species. However, across temperatures, C. flavipes yielded the highest number of offspring, followed by C. sesamiae and C. chilonis. The sex ratios did not vary significantly with species and temperature. Thus, the reproductive potentials of C. sesamiae and C. flavipes were greater than that of C. chilonis.

A viral lectin encoded in Cotesia plutellae bracovirus and its immunosuppressive effect on host hemocytes

An endoparasitoid wasp, Cotesia plutellae, induces immunosuppression of the host diamondback moth, Plutella xylostella. To identify an immunosuppressive factor, the parasitized hemolymph of P. xylostella was separated into plasma and hemocyte fractions. When nonparasitized hemocytes were overlaid with parasitized plasma, they showed significant reduction in bacterial binding efficacy. Here, we considered a viral lectin previously known in other Cotesia species as a humoral immunosuppressive candidate in C. plutellae parasitization. Based on consensus regions of the viral lectins, the corresponding lectin gene was cloned from P. xylostella parasitized by C. plutellae. Its cDNA is 674 bp long and encodes 157 amino acid residues containing a signal peptide (15 residues) and one carbohydrate recognition domain. Open reading frame is divided by one intron (156 bp) in its genomic DNA. Amino acid sequence shares 80% homology with that of C. ruficrus bracovirus lectin and is classified into C-type lectin. Southern hybridization analysis indicated that the cloned lectin gene was located at C. plutellae bracovirus (CpBV) genome. Both real-time quantitative RT-PCR and immunoblotting assays indicated that CpBV-lectin showed early expression during the parasitization. A recombinant CpBV-lectin was expressed in a bacterial system and the purified protein significantly inhibited the association between bacteria and hemocytes of nonparasitized P. xylostella. In the parasitized P. xylostella, CpBV-lectin was detected on the surface of parasitoid eggs after 24 h parasitization by its specific immunostaining. The 24 h old eggs were not encapsulated in vitro by hemocytes of P. xylostella, compared to newly laid parasitoid eggs showing no CpBV-lectin detectable and easily encapsulated. These results support an existence of a polydnaviral lectin family among Cotesia-associated bracovirus and propose its immunosuppressive function.