Suppressing effect of gamma-irradiated weevils on progeny production in the West Indian sweetpotato weevil Euscepes postfasciatus (Coleoptera: Curculionidae)

The West Indian sweet potato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) is a major pest of sweet potato Ipomoea batatas (L.) in the tropical and subtropical regions. The sterile insect technique (SIT) could be used as one of the most effective methods for suppression or eradication of the weevil. The effectiveness of SIT depends on the ability of the released sterile males to mate with and inseminate wild females. However, the effect of sterile weevils on the fitness of E. postfasciatus has not been evaluated on natural density. Here, we investigated the effect of gamma-irradiated weevil density on the number of weevil progeny. When irradiated weevils were released in numbers equal to those of non-irradiated weevils, the number of progeny was reduced by half of that in the control treatment, and it remained at this state for 2?weeks. Our results show that irradiated weevils ensure adequate and efficient suppression of wild weevils. We conclude that the SIT programs will be employed as effective eradication method for E. postfasciatus.     

Pleiotropic Impact of Endosymbiont Load and Co-Occurrence in the Maize Weevil Sitophilus zeamais

Individual traits vary among and within populations, and the co-occurrence of different endosymbiont species within a host may take place under varying endosymbiont loads in each individual host. This makes the recognition of the potential impact of such endosymbiont associations in insect species difficult, particularly in insect pest species. The maize weevil, Sitophilus zeamais Motsch. (Coleoptera: Curculionidae), a key pest species of stored cereal grains, exhibits associations with two endosymbiotic bacteria: the obligatory endosymbiont SZPE (“Sitophilus zeamais Primary Endosymbiont”) and the facultative endosymbiont Wolbachia. The impact of the lack of SZPE in maize weevil physiology is the impairment of nutrient acquisition and energy metabolism, while Wolbachia is an important factor in reproductive incompatibility. However, the role of endosymbiont load and co-occurrence in insect behavior, grain consumption, body mass and subsequent reproductive factors has not yet been explored. Here we report on the impacts of co-occurrence and varying endosymbiont loads achieved via thermal treatment and antibiotic provision via ingested water in the maize weevil. SZPE exhibited strong effects on respiration rate, grain consumption and weevil body mass, with observed effects on weevil behavior, particularly flight activity, and potential consequences for the management of this pest species. Wolbachia directly favored weevil fertility and exhibited only mild indirect effects, usually enhancing the SZPE effect. SZPE suppression delayed weevil emergence, which reduced the insect population growth rate, and the thermal inactivation of both symbionts prevented insect reproduction. Such findings are likely important for strain divergences reported in the maize weevil and their control, aspects still deserving future attention.     

Ultraviolet light-emitting diode (UV LED) trap the West Indian sweet potato weevil, <Emphasis Type="Italic">Euscepes postfasciatus</Emphasis> (Coleoptera: Curculionidae)

The West Indian sweet potato weevil Euscepes postfasciatus (Fairmaire) is a troublesome pest insect of sweet potato that originally came from the Caribbean, but is now expanding its distribution into the Pacific Islands. Although sterile insect techniques have been used against this pest in a demonstration experiment on Kume Island [Ohno et al. (2006) Kontyu to Shizen 41:25–30], effective methods of monitoring E. postfasciatus are scarce. It is necessary to detect the weevils at an early stage of invasion in uninvaded areas, and an attractant trap can be used to achieve this. Thus, we developed an ultraviolet (UV) light-emitting diode trap, invented a method for diffusing the light to attract more insects, and investigated the attractiveness of the light trap to E. postfasciatus under laboratory conditions. Our results indicate that diffused UV light has a higher potential to attract E. postfasciatus than direct UV light. Furthermore, sweet potato is an effective bait to use to capture the weevils attracted by UV light. Thus, E. postfasciatus can be trapped using diffused UV light and sweet potato bait.     

Appropriate number of inoculated eggs for mass-rearing the West Indian sweet potato weevil, <Emphasis Type="Italic">Euscepes postfasciatus</Emphasis> (Coleoptera: Curculionidae)

It is necessary to establish an economical mass-rearing system for the West Indian sweet potato weevil Euscepes postfasciatus (Fairmaire) so that large numbers can be created for use in the sterile insect technique (SIT) to control wild infestations. Both the quality of offspring and the number of adult weevils produced are important. As an adult female with a dry weight of less than 1.4 mg has an extremely low rate of production of offspring, we investigated the effect of an inoculated dose of egg suspension on both yield rate and body size, in order to determine the appropriate dose to attain the maximum yield of productive females. The number of emerged weevils increased as the inoculated dose of egg suspension increased. In brief, both yield rate and female body size significantly decreased as the inoculated dose of egg suspension increased, perhaps as a result of density effects. We conclude that the appropriate dose of inoculated egg suspension to attain the maximum yield from productive females of E. postfasciatus in the current mass-rearing system is 3 mL (containing approximately 450 eggs) per rearing container.     

Diversification of endosymbiosis: replacements,co-speciation and promiscuity of bacteriocyte symbionts in weevils

The processes and mechanisms underlying the diversification of host–microbe endosymbiotic associations are of evolutionary interest. Here we investigated the bacteriocyte-associated primary symbionts of weevils wherein the ancient symbiont Nardonella has experienced two independent replacement events: once by Curculioniphilus symbiont in the lineage of Curculio and allied weevils of the tribe Curculionini, and once by Sodalis-allied symbiont in the lineage of grain weevils of the genus Sitophilus. The Curculioniphilus symbiont was detected from 27 of 36 Curculionini species examined, the symbiont phylogeny was congruent with the host weevil phylogeny, and the symbiont gene sequences exhibited AT-biased nucleotide compositions and accelerated molecular evolution. These results suggest that the Curculioniphilus symbiont was acquired by an ancestor of the tribe Curculionini, replaced the original symbiont Nardonella, and has co-speciated with the host weevils over evolutionary time, but has been occasionally lost in several host lineages. By contrast, the Sodalis-allied symbiont of Sitophilus weevils exhibited no host–symbiont co-speciation, no AT-biased nucleotide compositions and only moderately accelerated molecular evolution. These results suggest that the Sodalis-allied symbiont was certainly acquired by an ancestor of the Sitophilus weevils and replaced the original Nardonella symbiont, but the symbiotic association must have experienced occasional re-associations such as new acquisitions, horizontal transfers, replacements and/or losses. We detected Sodalis-allied facultative symbionts in populations of the Curculionini weevils, which might represent potential evolutionary sources of the Sodalis-allied primary symbionts. Comparison of these newcomer bacteriocyte-associated symbiont lineages highlights potential evolutionary trajectories and consequences of novel symbionts after independent replacements of the same ancient symbiont.     

Effect of arbuscular mycorrhizal fungi (Glomus intraradices) on the oviposition of rice water weevil (Lissorhoptrus oryzophilus)

Root-feeding insects are important drivers in ecosystems, and links between aboveground oviposition preference and belowground larval performance have been suggested. The root-colonizing arbuscular mycorrhizal fungi (AMF) play a central role in plant nutrition and are known to change host quality for root-feeding insects. However, it is not known if and how AMF affect the aboveground oviposition of insects whose offspring feed on roots. According to the preference–performance hypothesis, insect herbivores oviposit on plants that will maximize offspring performance. In a greenhouse experiment with rice (Oryza sativa), we investigated the effects of AMF (Glomus intraradices) on aboveground oviposition of rice water weevil (Lissorhoptrus oryzophilus), the larvae of which feed belowground on the roots. Oviposition (i.e., the numbers of eggs laid by weevil females in leaf sheaths) was enhanced when the plants were colonized by AMF. However, the leaf area consumed by adult weevils was not affected. Although AMF reduced plant biomass, it increased nitrogen (N) and phosphorus concentrations in leaves and N in roots. The results suggest that rice water weevil females are able to discriminate plants for oviposition depending on their mycorrhizal status. The discrimination is probably related to AMF-mediated changes in plant quality, i.e., the females choose to oviposit more on plants with higher nutrient concentrations to potentially optimize offspring performance. AMF-mediated change in plant host choice for chewing insect oviposition is a novel aspect of below- and aboveground interactions.     

Induction of Furano-terpenoids in Sweet Potato Roots by the Larval Components of the Sweet Potato Weevils

When sweet potato root tissues were infested by the larvae of sweet potato weevil, Cylas formicarius and West Indian sweet potato weevil, Euscepes postfasciatus, furano-terpenoids and coumarins were produced in brown necrotic layer formed during the infestation.

The larval homogenates of both weevils also induced in the tissue the production of furano-terpenoids and coumarins, as well as the formation of necrotic layer. The larval homogenate of sweet potato weevil induced also ethylene formation, the marker of injury in the tissue. Investigations on the furano-terponoid inducing factor demonstrated that the factor was 20 mm KCl-soluble, non-dialyzable, acetone-precipitable, (NH₄)₂SO₄-precipitable, heat-unstable, passing through Sephadex G–25 column without sieving and partially inactivated by pronase, indicating that the factor was a high molecular weight compound, perhaps of a proteinacious property. It is likely that the factor causes injury or death to sweet potato root tissue, leading to the formation of ethylene and necrotic layer, and then to production of furano-terpenoids and coumarins.     

Gut microbiota of the pine weevil degrades conifer diterpenes and increases insect fitness

The pine weevil (Hylobius abietis), a major pest of conifer forests throughout Europe, feeds on the bark and cambium, tissues rich in terpenoid resins that are toxic to many insect herbivores. Here, we report the ability of the pine weevil gut microbiota to degrade the diterpene acids of Norway spruce. The diterpene acid levels present in ingested bark were substantially reduced on passage through the pine weevil gut. This reduction was significantly less upon antibiotic treatment, and supplementing the diet with gut suspensions from untreated insects restored the ability to degrade diterpenes. In addition, cultured bacteria isolated from pine weevil guts were shown to degrade a Norway spruce diterpene acid. In a metagenomic survey of the insect's bacterial community, we were able to annotate several genes of a previously described diterpene degradation (dit) gene cluster. Antibiotic treatment disrupted the core bacterial community of H. abietis guts and eliminated nearly all dit genes concordant with its reduction in diterpene degradation. Pine weevils reared on an artificial diet spiked with diterpenes, but without antibiotics, were found to lay more eggs with a higher hatching rate than weevils raised on diets with antibiotics or without diterpenes. These results suggest that gut symbionts contribute towards host fitness, but not by detoxification of diterpenes, as these compounds do not show toxic effects with or without antibiotics. Rather the ability to thrive in a terpene‐rich environment appears to allow gut microbes to benefit the weevil in other ways, such as increasing the nutritional properties of their diet.     

How the rice weevil breaks down the pectin network: Enzymatic synergism and sub-functionalization

Pectin is the most complex polysaccharide in nature and highly abundant in plant cell walls and middle lamellae, where it functions in plant growth and development. Phytopathogens utilize plant pectin as an energy source through enzyme-mediated degradation. These pectolytic enzymes include polygalacturonases (PGs) of the GH28 family and pectin methylesterases (PMEs) of the CE8 family. Recently, PGs were also identified in herbivorous insects of the distantly related plant bug, stick insect and Phytophaga beetle lineages. Unlike all other insects, weevils possess PMEs in addition to PGs. To investigate pectin digestion in insects and the role of PMEs in weevils, all PME and PG family members of the rice weevil Sitophilus oryzae were heterologously expressed and functionally characterized. Enzymatically active and inactive PG and PME family members were identified. The loss of activity can be explained by a lack of substrate binding correlating with substitutions of functionally important amino acid residues. We found subfunctionalization in both enzyme families, supported by expression pattern and substrate specificities as well as evidence for synergistic pectin breakdown. Our data suggest that the rice weevil might be able to use pectin as an energy source, and illustrates the potential of both PG and PME enzyme families to functionally diversify after horizontal gene transfer.     

What can a weevil teach a fly,and reciprocally? Interaction of host immune systems with endosymbionts in <Emphasis Type="Italic">Glossina</Emphasis> and <Emphasis Type="Italic">Sitophilus</Emphasis>

The tsetse fly (Glossina genus) is the main vector of African trypanosomes, which are protozoan parasites that cause human and animal African trypanosomiases in Sub-Saharan Africa. In the frame of the IAEA/FAO program ‘Enhancing Vector Refractoriness to Trypanosome Infection’, in addition to the tsetse, the cereal weevil Sitophilus has been introduced as a comparative system with regards to immune interactions with endosymbionts. The cereal weevil is an agricultural pest that destroys a significant proportion of cereal stocks worldwide. Tsetse flies are associated with three symbiotic bacteria, the multifunctional obligate Wigglesworthia glossinidia, the facultative commensal Sodalis glossinidius and the parasitic Wolbachia. Cereal weevils house an obligatory nutritional symbiosis with the bacterium Sodalis pierantonius, and occasionally Wolbachia. Studying insect host-symbiont interactions is highly relevant both for understanding the evolution of symbiosis and for envisioning novel pest control strategies. In both insects, the long co-evolution between host and endosymbiont has led to a stringent integration of the host-bacteria partnership. These associations were facilitated by the development of specialized host traits, including symbiont-housing cells called bacteriocytes and specific immune features that enable both tolerance and control of the bacteria. In this review, we compare the tsetse and weevil model systems and compile the latest research findings regarding their biological and ecological similarities, how the immune system controls endosymbiont load and location, and how host-symbiont interactions impact developmental features including cuticle synthesis and immune system maturation. We focus mainly on the interactions between the obligate symbionts and their host’s immune systems, a central theme in both model systems. Finally, we highlight how parallel studies on cereal weevils and tsetse flies led to mutual discoveries and stimulated research on each model, creating a pivotal example of scientific improvement through comparison between relatively distant models.     

The Facultative Symbiont Rickettsia Protects an Invasive Whitefly against Entomopathogenic Pseudomonas syringae Strains

Facultative endosymbionts can benefit insect hosts in a variety of ways, including context-dependent roles, such as providing defense against pathogens. The role of some symbionts in defense may be overlooked, however, when pathogen infection is transient, sporadic, or asymptomatic. The facultative endosymbiont Rickettsia increases the fitness of the sweet potato whitefly (Bemisia tabaci) in some populations through mechanisms that are not yet understood. In this study, we investigated the role of Rickettsia in mediating the interaction between the sweet potato whitefly and Pseudomonas syringae, a common environmental bacterium, some strains of which are pathogenic to aphids. Our results show that P. syringae multiplies within whiteflies, leading to host death, and that whiteflies infected with Rickettsia show a decreased rate of death due to P. syringae. Experiments using plants coated with P. syringae confirmed that whiteflies can acquire the bacteria at a low rate while feeding, leading to increased mortality, particularly when the whiteflies are not infected with Rickettsia. These results suggest that P. syringae may affect whitefly populations in nature and that Rickettsia can ameliorate this effect. This study highlights the possible importance of interactions among opportunistic environmental pathogens and endosymbionts of insects.     

Evaluation and implications of Andean potato weevil infestation sources for its management in the Andean region

The Andean potato weevil Premnotrypes suturicallus Kuschel (Coleoptera: Curculionidae) is one of the most damaging potato (Solanum spp.) pests in the Andes. The objective of this study was to get a better understanding of weevil infestation sources and its distribution in potato fields as well as on the effect of potato cropping intensity and farmers’ harvest practices on weevil infestation to provide better clues for its management. For this purpose, a methodology was developed to assess weevil field densities in potato. A Taylor’s power law indicated that 177 and 69 samples are necessary to assess low (0.5 weevils/m²) and high (5 weevils/m²) weevil densities for a moderate reliability or precision level (D = 0.5). In potato fields, overwintering weevil densities were highest at field borders with 3.5 weevils/m² at a distance of <2.5 m compared to 0.7 weevils/m² at distances >10 m from the field border. No effects of time of harvest could be detected on soil overwintering weevil densities. The quantification of the larval density per potato plant after harvest showed that fields after 2‐year potato rotations had eight times more overwintering weevils compared to 1‐year rotations. Weevil infestation sources were mainly potato fields with the highest infestation (89%) followed by olluco (Ullucus tuberosus) and oat (Avena sativa) fields having volunteer potato plants (35%). The results confirm and support previous assumptions about the importance of the cropping systems for Andean potato weevil infestation and management. The confirmation that weevils do not occur or only in neglected numbers on fallow fields supports the use of plastic barriers to effectively exclude migrating flightless adult weevils to potato fields cultivated after fallow. The distribution of overwintering weevils indicates that farmers could concentrate efforts to control adult weevils mainly to the first meters of potato fields.     

The neogregarine protozoan Farinocystis sp. reduces longevity and fecundity in the West Indian sweet potato weevil, Euscepes postfasciatus (Fairmaire)

The number of West Indian sweet potato weevils, Euscepes postfasciatus, being mass-reared in a facility for use in sterile insect technique (SIT) eradication programs has undergone a drastic reduction. A neogregarine protozoan pathogen Farinocystis sp. (an undescribed species) was detected in vivo in the mass-reared E. postfasciatus. We investigated the effects of this disease on the longevity and fecundity of host weevils and the incubation time of the disease in the host body under mass-rearing conditions. Our results demonstrated that infection by this Farinocystis sp. decreased both longevity and fecundity in E. postfasciatus. In particular, the pathogen severely limited the production of progeny by infected females compared to healthy females. Therefore, we consider this protozoan infection to be the major cause of the decreased E. postfasciatus production in the mass-rearing facility.     

Reduced dispersal and survival in the sweet potato weevil (Euscepes postfasciatus) after irradiation

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Sublethal effects of malathion on boll weevil (Coleoptera: Curculionidae) fecundity when maintained on cotton squares or artificial diet

Mated 3‐day‐old female boll weevils, Anthonomus grandis grandis Boheman, reared from field‐infested cotton (Gossypium hirsutum L.) squares were topically treated with an estimated LD₅₀ of malathion (2 μg) to assess its effects on fecundity, oviposition, and body fat condition. Two different food sources, cotton squares and artificial diet, were assessed in malathion‐treated and nontreated (control) weevils. The LD₅₀ caused ～50% mortality in the square‐fed malathion treatment, but the artificial diet‐fed malathion‐treated weevils were less susceptible. LD₅₀ survivors fed on the squares produced ≥ 9 times more chorionated eggs in the ovaries and oviposited ≥ 19‐fold more than survivors fed artificial diet, regardless of the malathion treatment. Boll weevils that survived a 2 μg LD₅₀ malathion and also fed squares were ～4.5‐fold leaner than diet‐fed weevils. Our findings demonstrate that non‐resistant boll weevils surviving a sublethal dose of malathion will reproduce without any delay or significant loss in fecundity, and the food source for which boll weevils are maintained when conducting these assays will directly affect the results. The significance of these findings and how they are related to the final stages of eradicating the boll weevil from the US are discussed.     

Mutual benefit interactions between banded pine weevil Pissodes castaneus and blue‐stain fungus Leptographium serpens in maritime pine

• 1 Ecological interactions between banded pine weevil Pissodes castaneus and blue‐stain fungus Leptographium serpens, when simultaneously sharing the same host plant (maritime pine Pinus pinaster) in winter and spring, were investigated. Temporal components of the interaction were taken into account by either introducing the weevils and the pathogen simultaneously or sequentially, with the weevils being introduced 1 month after the fungal inoculation.
• 2 We measured larval mortality, development time, offspring number, sex ratio and body size of P. castaneus. Phloem phosphorus and nitrogen concentrations were also assessed. Furthermore, we tested whether: (i) emerging offspring transported propagules of the fungus; (ii) artificially‐contaminated weevils may transmit the disease to healthy trees; and (iii) field collected P. castaneus carry the fungus.
• 3 The fungus enhanced weevil colonization and brood production in both seasons. During winter and spring, adults from trees where the pathogen was inoculated prior to weevil introduction emerged earlier than weevils from trees where they had been introduced simultaneously with the fungus. During winter, weevils from pre‐inoculated trees were also larger. Sex ratio and larval mortality were not affected. Leptographium serpens did not affect phloem nitrogen content but phosphorus content was greater in plants inoculated with the pathogen, which may explain the findings on weevil growth.
• 4 Sixty‐five percent of the weevils that emerged from inoculated trees carried spores of L. serpens, although no successful isolation was made from field collected weevils. The fungus was recovered from 25% of the trees infested with artificially‐contaminated weevils.
• 5 These results suggest that P. castaneus benefits from the presence of L. serpens and may contribute to its spread.

     

What can a weevil teach a fly,and reciprocally? Interaction of host immune systems with endosymbionts in Glossina and Sitophilus

The tsetse fly (Glossina genus) is the main vector of African trypanosomes, which are protozoan parasites that cause human and animal African trypanosomiases in Sub-Saharan Africa. In the frame of the IAEA/FAO program ‘Enhancing Vector Refractoriness to Trypanosome Infection’, in addition to the tsetse, the cereal weevil Sitophilus has been introduced as a comparative system with regards to immune interactions with endosymbionts. The cereal weevil is an agricultural pest that destroys a significant proportion of cereal stocks worldwide. Tsetse flies are associated with three symbiotic bacteria, the multifunctional obligate Wigglesworthia glossinidia, the facultative commensal Sodalis glossinidius and the parasitic Wolbachia. Cereal weevils house an obligatory nutritional symbiosis with the bacterium Sodalis pierantonius, and occasionally Wolbachia. Studying insect host-symbiont interactions is highly relevant both for understanding the evolution of symbiosis and for envisioning novel pest control strategies. In both insects, the long co-evolution between host and endosymbiont has led to a stringent integration of the host-bacteria partnership. These associations were facilitated by the development of specialized host traits, including symbiont-housing cells called bacteriocytes and specific immune features that enable both tolerance and control of the bacteria. In this review, we compare the tsetse and weevil model systems and compile the latest research findings regarding their biological and ecological similarities, how the immune system controls endosymbiont load and location, and how host-symbiont interactions impact developmental features including cuticle synthesis and immune system maturation. We focus mainly on the interactions between the obligate symbionts and their host’s immune systems, a central theme in both model systems. Finally, we highlight how parallel studies on cereal weevils and tsetse flies led to mutual discoveries and stimulated research on each model, creating a pivotal example of scientific improvement through comparison between relatively distant models.

     

Effect of temperature on the development of the West Indian sweet potato weevil, <Emphasis Type="Italic">Euscepes postfasciatus</Emphasis> (Fairmaire) (Coleoptera: Curculionidae) on an artificial diet

The relationship between temperature and the development of the West Indian sweet potato weevil, Euscepes postfasciatus, on an artificial larval diet containing powdered sweet potato root, was examined at different fixed temperatures from 22 to 31°C. The developmental periods for egg, larvae, and pupae stages shortened in correlation with increased temperature. The thermal constant was 769.2 degree-days and the developmental zero for female and male was 11.1 and 11.7°C, respectively. Although we can rear this weevil at temperatures ranging from 22 to 31°C, rearing temperatures should be kept between 25 and 28°C because the developmental stages were too long at 22°C and the larval period was delayed at 31°C. The basis for these developmental data will be a useful key factor in designing a plan to eradicate the weevil by using a mass-rearing system and SIT.     

Field release of a prospective biological control agent of weeds,Ceratapion basicorne,to evaluate potential risk to a nontarget crop

Ceratapion basicorne is a prospective biological control agent of yellow starthistle (Centaurea solstitialis), which is an invasive alien weed in the USA. Although the weevil has a strong preference for yellow starthistle, it has been reported to develop sometimes on safflower in larval transfer and no-choice experiments. Although safflower was not attacked by this insect in previous field experiments, a release permit was denied because of concern for risk to safflower. Adult C. basicorne were released in a field experiment in which two varieties of safflower were grown in solid blocks on either side of a small number of yellow starthistle plants. Plants were dissected at the time of weevil pupation. Immature insects were reared to adult stage on artificial diet or were preserved in acetone to identify by molecular genetic analysis. C. basicorne infested 54% of the yellow starthistle plants and 0% of 1021 safflower plants. A different weevil, Ceratapion orientale, infested 1.5% of the safflower plants. These results corroborate two other published field studies in which C. basicorne was not reared from safflower. The combined results of nine experiments provide a point estimate that the probability of attack is less than 0.00059, with 99.9% confidence that it is less than 0.0045. The consistency of results from field experiments in three countries and the absence of any report of this insect being reared from safflower in the field in the weevil’s native range support the conclusion that this insect poses no significant risk to safflower.