Electrophoretic variation at three enzyme loci‐alcohol dehydrogenase (Adh), glycerophosphate dehy... more Electrophoretic variation at three enzyme loci‐alcohol dehydrogenase (Adh), glycerophosphate dehydrogenase (Gpdh), triosephosphate isomerase (Tpi)‐ is compared in Australian Drosophila melanogaster populations at three levels of spatial heterogeneity; among breeding sites, within populations, and between populations at the geographic level. Heterogeneity at the breeding site level greatly exceeds that among adults within populations, indicating greater intermixing at the mobile adult stage than at the developmentally immature and less migratory larval stage. Heterogeneity at the microspatial level is large relative to the geographic level at two of these loci.Spatial patterns of variation in ecological phenotypes are also considered. It is argued that electrophoretic variants may contribute little to an understanding of this quantitative variation, and that a more useful approach in ecological genetics is to consider ecological phenotypes as primary data.
Comparative analyses of ectotherm susceptibility to climate change often focus on thermal extreme... more Comparative analyses of ectotherm susceptibility to climate change often focus on thermal extremes, yet responses to aridity may be equally important. Here we focus on plasticity in desiccation resistance, a key trait shaping distributions of <i>Drosophila</i> species and other small ectotherms. We examined the extent to which 32 <i>Drosophila</i> species, varying in their distribution, could increase their desiccation resistance via phenotypic plasticity involving hardening, linking these responses to environment, phylogeny and basal resistance. We found no evidence to support the seasonality hypothesis; species with higher hardening plasticity did not occupy environments with higher and more seasonal precipitation. As basal resistance increased, the capacity of species to respond via phenotypic plasticity decreased, suggesting plastic responses involving hardening may be constrained by basal resistance. Trade-offs between basal desiccation resistance and plasticity were not universal across the phylogeny and tended to occur within specific clades. Phylogeny, environment and trade-offs all helped to explain variation in plasticity for desiccation resistance but in complex ways. These findings suggest some species have the ability to counter dry periods through plastic responses, whereas others do not; and this ability will depend to some extent on a species' placement within a phylogeny, along with its basal level of resistance.
Wolbachia bacteria are being widely released for suppression of dengue transmitted by Aedes mosqu... more Wolbachia bacteria are being widely released for suppression of dengue transmitted by Aedes mosquitoes. Walker, Quek, Jeffries and colleagues present robust evidence for natural Wolbachia infections in malaria-vectoring Anopheles mosquitoes, paving the way for new Wolbachia-based interventions.
Spodoptera frugiperda has been reported in China since the beginning of 2019, threatening maize p... more Spodoptera frugiperda has been reported in China since the beginning of 2019, threatening maize production and raising questions about interactions with indigenous pests on maize. Spodoptera frugiperda is in the same feeding guild as the indigenous species Ostrinia furnacalis , with both pests attacking maize plant tissues such as whorls and tassels. In this study, the interaction of these species was assessed under laboratory and field conditions by examining the survival and predation rate of larvae in maize whorls and tassels. Intraspecific and interspecific interactions of larvae were assessed to characterize movements in arenas in the presence and absence of food. When S. frugiperda and O. furnacalis were present at an equal ratio, S. frugiperda prevailed over O. furnacalis with a high survival rate (over 90%) and predation rate of O. furnacalis (over 40%) under most scenarios in both laboratory and field conditions. When both species occurred at different ratios in the field, S. frugiperda had on average almost 16 times the number of survivors as O. furnacalis on whorls, and 8.3 times the number on tassels, highlighting a substantial advantage in these environments. Spodoptera frugiperda larvae undertook a low frequency of lethal attacks (strikes) and showed a higher incidence of defensive movements compared to O. furnacalis . A field survey showed that S. frugiperda larvae can colonize vegetative plant stages in maize, but it remains to be seen if competitive interactions can be linked to population dynamics of both species though they are expected to favor S. frugiperda .
Background: Wolbachia wMel is the most commonly used strain in rear and release strategies for Ae... more Background: Wolbachia wMel is the most commonly used strain in rear and release strategies for Aedes aegypti mosquitoes that aim to inhibit the transmission of arboviruses such as dengue, Zika, Chikungunya and yellow fever. However, the long-term establishment of wMel in natural Ae. aegypti populations raises concerns that interactions between Wolbachia wMel and Ae. aegypti may lead to changes in the host genome, which could affect useful attributes of Wolbachia that allow it to invade and suppress disease transmission. Results: We applied an evolve-and-resequence approach to study genome-wide genetic changes in Ae. aegypti from the Cairns region, Australia, where Wolbachia wMel was first introduced more than 10 years ago. Mosquito samples were collected at three different time points in Gordonvale, Australia, covering the phase before (2010) and after (2013 and 2018) Wolbachia releases. An additional three locations where Wolbachia replacement happened at different times across the last decade were also sampled in 2018. We found that the genomes of mosquito populations mostly remained stable after Wolbachia release, with population differences tending to reflect the geographic location of the populations rather than Wolbachia infection status. However, outlier analysis suggests that Wolbachia may have had an influence on some genes related to immune response, development, recognition and behavior. Conclusions: Ae. aegypti populations remained geographically distinct after Wolbachia wMel releases in North Australia despite their Wolbachia infection status. At some specific genomic loci, we found signs of selection associated with Wolbachia, suggesting potential evolutionary impacts can happen in the future and further monitoring is warranted.
We have started to test the effects of environmental extremes on the expression of genetic variat... more We have started to test the effects of environmental extremes on the expression of genetic variation for traits likely to be under selection in natural populations. We have shown that field heritability may be high for stress response traits in contrast to morphological traits, which tend to show lower levels of heritable variation in nature compared with the laboratory. Selection for increased stress resistance can lead to a number of other evolutionary changes, and these may underlie trade-offs between favourable and stressful environments. Temperature extremes can have a marked influence on the heritability of life history traits. Heritabilities for fecundity can be high when parental flies are reared at low temperatures and under field conditions. The expression of genetic variation for development time is somewhat more complex when temperature extremes are considered. Populations at species margins may be ideal for studying the effects of environmental stress on evolution.
Highlights d The wStri Wolbachia infection was stably established in Nilaparvata lugens (Stå l) d... more Highlights d The wStri Wolbachia infection was stably established in Nilaparvata lugens (Stå l) d Extremely high density of wStri Wolbachia was observed in all examined tissues d High CI level ensures rapid spread of wStri Wolbachia in laboratory populations d wStri Wolbachia blocks rice virus transmission involving vector Nilaparvata lugens
Aedes notoscriptus (Skuse) is a container-inhabiting mosquito endemic to Australia that vectors a... more Aedes notoscriptus (Skuse) is a container-inhabiting mosquito endemic to Australia that vectors arboviruses and is suspected to transmit Mycobacterium ulcerans, the cause of Buruli ulcer. We evaluated the effectiveness of the In2Care station, which suppresses mosquito populations via the entomopathogenic fungus, Beauveria bassiana, and the insect growth regulator pyriproxyfen, the latter of which is autodisseminated among larval habitats by contaminated mosquitoes. A field trial was conducted using 110 In2Care stations in a 50,000 m 2 area and results were compared to 4 control areas that did not receive the treatment. Efficacy was evaluated by comparing egg counts and measuring larvicidal impact in surrounding breeding sites. Laboratory experiments validated the effect of B. bassiana on adult survival. Results of this field trial indicate that, 6 wk after the In2Care stations were deployed, treatment site ovitraps contained 43% fewer eggs than control site ovitraps, and 33% fewer eggs after 10 wk, suggesting that the In2Care station was able to reduce the egg density of Ae. notoscriptus. Population reduction remained evident for up to 3 wk after In2Care stations were removed. Treatment site ovitraps had significantly fewer Ae. notoscriptus eclosing than control site ovitraps, confirming the pyriproxyfen autodissemination feature of the stations. An average reduction of 50% in adult eclosion was achieved. Exposure to B. bassiana resulted in four-times higher mortality among adult mosquitoes. Additionally, using fresh In2Care nettings led to an 88% decrease in average survival compared to 4-wk-old nettings. The use of In2Care stations has potential for suppressing Ae. notoscriptus egg density.
Colonizing populations of alien invasive species are often unstable due to their small population... more Colonizing populations of alien invasive species are often unstable due to their small population size, ongoing gene ow, and ongoing adaptation to local conditions. These processes should lead to molecular signatures at the population level. However, temporal changes in genetic patterns after introduction are rarely examined. The western ower thrips (WFT), Frankliniella occidentalis (Pergande), is a globally invasive pest of vegetables and ornamental crops. Early colonized populations in China exhibited a strong population structure re ecting different invasion sources. Here, we reexamine this pattern after ten years. Over this period, the number of alleles has declined signi cantly and strong population genetic structure found early in the invasion stage has been lost. A high level of gene ow between both geographically close and distant populations was identi ed. The reduced allele number and loss of population structure might re ect both ongoing gene ow and ongoing control measures to suppress populations. Our results emphasize the importance of continuing to manage gene ow among invaded areas to limit the exchange of alleles that might facilitate further adaptive changes following invasion.
The Scaptodrosophila genus represents a large group of drosophilids with a worldwide distribution... more The Scaptodrosophila genus represents a large group of drosophilids with a worldwide distribution and a predominance of species in Australia, but there is little information on the presence and impacts of Wolbachia endosymbionts in this group. Here we describe the first Wolbachia infection from this group, wClay isolated from Scaptodrosophila claytoni (van Klinken), a species from the east coast of Australia. The infection is polymorphic in natural populations, occurring at a frequency of around 6%–10%. wClay causes male killing, producing female‐biased lines; most lines showed 100% male killing, though in 1 line it was &lt;80%. The lines need to be maintained through the introduction of males unless the infection is removed by tetracycline treatment. wClay is transmitted at a high fidelity (98.6%) through the maternal lineage and has been stable in 2 laboratory lines across 24 generations, suggesting it is likely to persist in populations. The infection has not been previously described but is closely related to the male‐killing Wolbachia recently described from Drosophila pandora based on multilocus sequence typing and the wsp gene. Male‐killing Wolbachia are likely to be common in drosophilids but remain difficult to detect because the infections can often be at a low frequency.
Macroinvertebrate biodiversity is routinely used to assess the ecological condition of freshwater... more Macroinvertebrate biodiversity is routinely used to assess the ecological condition of freshwater environments. Macroinvertebrates are traditionally identified morphologically to mostly family level but using DNA metabarcoding they can be rapidly and reliably identified to species. Developing standardised, robust and cost-effective protocols would enable DNA metabarcoding to be broadly used for routine freshwater biological assessments leading to both potential cost savings as well as increased taxonomic resolution. To further reduce the cost and time it takes to process samples, we examine the feasibility of DNA metabarcoding unsorted macroinvertebrates (macroinvertebrates and debris) from sweep net samples collected from stream pool or edge habitats. We processed these unsorted samples with a standardised method and tested multiple primer sets widely used for invertebrate DNA metabarcoding. We found that currently available DNA metabarcoding primers are well suited for processing unsorted samples, though some performed better than others. However, macroinvertebrate density affected the number of species detected with DNA metabarcoding, particularly the detection of rare taxa. These findings show that DNA metabarcoding of unsorted net contents could streamline macroinvertebrate sample processing for bioassessment, but the optimal amounts of unsorted material for sub-sampling needs to be considered when assessing macroinvertebrate biodiversity. Hosted file Bulk sample paper_9_Oct_20_2.pdf available at https://authorea.com/users/367412/articles/ 486850-using-unsorted-sweep-net-samples-to-rapidly-assess-macroinvertebrate-biodiversity
While facultative endosymbionts are now known to protect insect hosts against pathogens and paras... more While facultative endosymbionts are now known to protect insect hosts against pathogens and parasitoids, the effects of endosymbionts on insecticide resistance are still unclear. Here we show that Wolbachia are associated with increased resistance to the commonly used insecticide, buprofezin, in the small brown planthopper (Laodelphax striatellus) in some genetic backgrounds while having no effect in other backgrounds. In three Wolbachia-infected lines from experimental buprofezin-resistant strains and one line from a buprofezin-susceptible line established from Chuxiong, Yunnan province, China, susceptibility to buprofezin increased after removal of Wolbachia. An increase in susceptibility was also evident in a Wolbachia-infected line established from a field population in Rugao, Jiangsu province. However, no increase was evident in two field populations from Nanjing and Fengxian, Jiangsu province, China. When Wolbachia was introgressed into different genetic backgrounds, followed by Wolbachia removal, the data pointed to Wolbachia effects that depend on the nuclear background as well as on the Wolbachia strain. However, there was no relationship between Wolbachia density and the component of buprofezin resistance associated with This article is protected by copyright. All rights reserved. 4 the symbiont. The results suggest that Wolbachia effects associated with chemical resistance are complex and unpredictable, but also that they can be substantial.
Electrophoretic variation at three enzyme loci‐alcohol dehydrogenase (Adh), glycerophosphate dehy... more Electrophoretic variation at three enzyme loci‐alcohol dehydrogenase (Adh), glycerophosphate dehydrogenase (Gpdh), triosephosphate isomerase (Tpi)‐ is compared in Australian Drosophila melanogaster populations at three levels of spatial heterogeneity; among breeding sites, within populations, and between populations at the geographic level. Heterogeneity at the breeding site level greatly exceeds that among adults within populations, indicating greater intermixing at the mobile adult stage than at the developmentally immature and less migratory larval stage. Heterogeneity at the microspatial level is large relative to the geographic level at two of these loci.Spatial patterns of variation in ecological phenotypes are also considered. It is argued that electrophoretic variants may contribute little to an understanding of this quantitative variation, and that a more useful approach in ecological genetics is to consider ecological phenotypes as primary data.
Comparative analyses of ectotherm susceptibility to climate change often focus on thermal extreme... more Comparative analyses of ectotherm susceptibility to climate change often focus on thermal extremes, yet responses to aridity may be equally important. Here we focus on plasticity in desiccation resistance, a key trait shaping distributions of <i>Drosophila</i> species and other small ectotherms. We examined the extent to which 32 <i>Drosophila</i> species, varying in their distribution, could increase their desiccation resistance via phenotypic plasticity involving hardening, linking these responses to environment, phylogeny and basal resistance. We found no evidence to support the seasonality hypothesis; species with higher hardening plasticity did not occupy environments with higher and more seasonal precipitation. As basal resistance increased, the capacity of species to respond via phenotypic plasticity decreased, suggesting plastic responses involving hardening may be constrained by basal resistance. Trade-offs between basal desiccation resistance and plasticity were not universal across the phylogeny and tended to occur within specific clades. Phylogeny, environment and trade-offs all helped to explain variation in plasticity for desiccation resistance but in complex ways. These findings suggest some species have the ability to counter dry periods through plastic responses, whereas others do not; and this ability will depend to some extent on a species' placement within a phylogeny, along with its basal level of resistance.
Wolbachia bacteria are being widely released for suppression of dengue transmitted by Aedes mosqu... more Wolbachia bacteria are being widely released for suppression of dengue transmitted by Aedes mosquitoes. Walker, Quek, Jeffries and colleagues present robust evidence for natural Wolbachia infections in malaria-vectoring Anopheles mosquitoes, paving the way for new Wolbachia-based interventions.
Spodoptera frugiperda has been reported in China since the beginning of 2019, threatening maize p... more Spodoptera frugiperda has been reported in China since the beginning of 2019, threatening maize production and raising questions about interactions with indigenous pests on maize. Spodoptera frugiperda is in the same feeding guild as the indigenous species Ostrinia furnacalis , with both pests attacking maize plant tissues such as whorls and tassels. In this study, the interaction of these species was assessed under laboratory and field conditions by examining the survival and predation rate of larvae in maize whorls and tassels. Intraspecific and interspecific interactions of larvae were assessed to characterize movements in arenas in the presence and absence of food. When S. frugiperda and O. furnacalis were present at an equal ratio, S. frugiperda prevailed over O. furnacalis with a high survival rate (over 90%) and predation rate of O. furnacalis (over 40%) under most scenarios in both laboratory and field conditions. When both species occurred at different ratios in the field, S. frugiperda had on average almost 16 times the number of survivors as O. furnacalis on whorls, and 8.3 times the number on tassels, highlighting a substantial advantage in these environments. Spodoptera frugiperda larvae undertook a low frequency of lethal attacks (strikes) and showed a higher incidence of defensive movements compared to O. furnacalis . A field survey showed that S. frugiperda larvae can colonize vegetative plant stages in maize, but it remains to be seen if competitive interactions can be linked to population dynamics of both species though they are expected to favor S. frugiperda .
Background: Wolbachia wMel is the most commonly used strain in rear and release strategies for Ae... more Background: Wolbachia wMel is the most commonly used strain in rear and release strategies for Aedes aegypti mosquitoes that aim to inhibit the transmission of arboviruses such as dengue, Zika, Chikungunya and yellow fever. However, the long-term establishment of wMel in natural Ae. aegypti populations raises concerns that interactions between Wolbachia wMel and Ae. aegypti may lead to changes in the host genome, which could affect useful attributes of Wolbachia that allow it to invade and suppress disease transmission. Results: We applied an evolve-and-resequence approach to study genome-wide genetic changes in Ae. aegypti from the Cairns region, Australia, where Wolbachia wMel was first introduced more than 10 years ago. Mosquito samples were collected at three different time points in Gordonvale, Australia, covering the phase before (2010) and after (2013 and 2018) Wolbachia releases. An additional three locations where Wolbachia replacement happened at different times across the last decade were also sampled in 2018. We found that the genomes of mosquito populations mostly remained stable after Wolbachia release, with population differences tending to reflect the geographic location of the populations rather than Wolbachia infection status. However, outlier analysis suggests that Wolbachia may have had an influence on some genes related to immune response, development, recognition and behavior. Conclusions: Ae. aegypti populations remained geographically distinct after Wolbachia wMel releases in North Australia despite their Wolbachia infection status. At some specific genomic loci, we found signs of selection associated with Wolbachia, suggesting potential evolutionary impacts can happen in the future and further monitoring is warranted.
We have started to test the effects of environmental extremes on the expression of genetic variat... more We have started to test the effects of environmental extremes on the expression of genetic variation for traits likely to be under selection in natural populations. We have shown that field heritability may be high for stress response traits in contrast to morphological traits, which tend to show lower levels of heritable variation in nature compared with the laboratory. Selection for increased stress resistance can lead to a number of other evolutionary changes, and these may underlie trade-offs between favourable and stressful environments. Temperature extremes can have a marked influence on the heritability of life history traits. Heritabilities for fecundity can be high when parental flies are reared at low temperatures and under field conditions. The expression of genetic variation for development time is somewhat more complex when temperature extremes are considered. Populations at species margins may be ideal for studying the effects of environmental stress on evolution.
Highlights d The wStri Wolbachia infection was stably established in Nilaparvata lugens (Stå l) d... more Highlights d The wStri Wolbachia infection was stably established in Nilaparvata lugens (Stå l) d Extremely high density of wStri Wolbachia was observed in all examined tissues d High CI level ensures rapid spread of wStri Wolbachia in laboratory populations d wStri Wolbachia blocks rice virus transmission involving vector Nilaparvata lugens
Aedes notoscriptus (Skuse) is a container-inhabiting mosquito endemic to Australia that vectors a... more Aedes notoscriptus (Skuse) is a container-inhabiting mosquito endemic to Australia that vectors arboviruses and is suspected to transmit Mycobacterium ulcerans, the cause of Buruli ulcer. We evaluated the effectiveness of the In2Care station, which suppresses mosquito populations via the entomopathogenic fungus, Beauveria bassiana, and the insect growth regulator pyriproxyfen, the latter of which is autodisseminated among larval habitats by contaminated mosquitoes. A field trial was conducted using 110 In2Care stations in a 50,000 m 2 area and results were compared to 4 control areas that did not receive the treatment. Efficacy was evaluated by comparing egg counts and measuring larvicidal impact in surrounding breeding sites. Laboratory experiments validated the effect of B. bassiana on adult survival. Results of this field trial indicate that, 6 wk after the In2Care stations were deployed, treatment site ovitraps contained 43% fewer eggs than control site ovitraps, and 33% fewer eggs after 10 wk, suggesting that the In2Care station was able to reduce the egg density of Ae. notoscriptus. Population reduction remained evident for up to 3 wk after In2Care stations were removed. Treatment site ovitraps had significantly fewer Ae. notoscriptus eclosing than control site ovitraps, confirming the pyriproxyfen autodissemination feature of the stations. An average reduction of 50% in adult eclosion was achieved. Exposure to B. bassiana resulted in four-times higher mortality among adult mosquitoes. Additionally, using fresh In2Care nettings led to an 88% decrease in average survival compared to 4-wk-old nettings. The use of In2Care stations has potential for suppressing Ae. notoscriptus egg density.
Colonizing populations of alien invasive species are often unstable due to their small population... more Colonizing populations of alien invasive species are often unstable due to their small population size, ongoing gene ow, and ongoing adaptation to local conditions. These processes should lead to molecular signatures at the population level. However, temporal changes in genetic patterns after introduction are rarely examined. The western ower thrips (WFT), Frankliniella occidentalis (Pergande), is a globally invasive pest of vegetables and ornamental crops. Early colonized populations in China exhibited a strong population structure re ecting different invasion sources. Here, we reexamine this pattern after ten years. Over this period, the number of alleles has declined signi cantly and strong population genetic structure found early in the invasion stage has been lost. A high level of gene ow between both geographically close and distant populations was identi ed. The reduced allele number and loss of population structure might re ect both ongoing gene ow and ongoing control measures to suppress populations. Our results emphasize the importance of continuing to manage gene ow among invaded areas to limit the exchange of alleles that might facilitate further adaptive changes following invasion.
The Scaptodrosophila genus represents a large group of drosophilids with a worldwide distribution... more The Scaptodrosophila genus represents a large group of drosophilids with a worldwide distribution and a predominance of species in Australia, but there is little information on the presence and impacts of Wolbachia endosymbionts in this group. Here we describe the first Wolbachia infection from this group, wClay isolated from Scaptodrosophila claytoni (van Klinken), a species from the east coast of Australia. The infection is polymorphic in natural populations, occurring at a frequency of around 6%–10%. wClay causes male killing, producing female‐biased lines; most lines showed 100% male killing, though in 1 line it was &lt;80%. The lines need to be maintained through the introduction of males unless the infection is removed by tetracycline treatment. wClay is transmitted at a high fidelity (98.6%) through the maternal lineage and has been stable in 2 laboratory lines across 24 generations, suggesting it is likely to persist in populations. The infection has not been previously described but is closely related to the male‐killing Wolbachia recently described from Drosophila pandora based on multilocus sequence typing and the wsp gene. Male‐killing Wolbachia are likely to be common in drosophilids but remain difficult to detect because the infections can often be at a low frequency.
Macroinvertebrate biodiversity is routinely used to assess the ecological condition of freshwater... more Macroinvertebrate biodiversity is routinely used to assess the ecological condition of freshwater environments. Macroinvertebrates are traditionally identified morphologically to mostly family level but using DNA metabarcoding they can be rapidly and reliably identified to species. Developing standardised, robust and cost-effective protocols would enable DNA metabarcoding to be broadly used for routine freshwater biological assessments leading to both potential cost savings as well as increased taxonomic resolution. To further reduce the cost and time it takes to process samples, we examine the feasibility of DNA metabarcoding unsorted macroinvertebrates (macroinvertebrates and debris) from sweep net samples collected from stream pool or edge habitats. We processed these unsorted samples with a standardised method and tested multiple primer sets widely used for invertebrate DNA metabarcoding. We found that currently available DNA metabarcoding primers are well suited for processing unsorted samples, though some performed better than others. However, macroinvertebrate density affected the number of species detected with DNA metabarcoding, particularly the detection of rare taxa. These findings show that DNA metabarcoding of unsorted net contents could streamline macroinvertebrate sample processing for bioassessment, but the optimal amounts of unsorted material for sub-sampling needs to be considered when assessing macroinvertebrate biodiversity. Hosted file Bulk sample paper_9_Oct_20_2.pdf available at https://authorea.com/users/367412/articles/ 486850-using-unsorted-sweep-net-samples-to-rapidly-assess-macroinvertebrate-biodiversity
While facultative endosymbionts are now known to protect insect hosts against pathogens and paras... more While facultative endosymbionts are now known to protect insect hosts against pathogens and parasitoids, the effects of endosymbionts on insecticide resistance are still unclear. Here we show that Wolbachia are associated with increased resistance to the commonly used insecticide, buprofezin, in the small brown planthopper (Laodelphax striatellus) in some genetic backgrounds while having no effect in other backgrounds. In three Wolbachia-infected lines from experimental buprofezin-resistant strains and one line from a buprofezin-susceptible line established from Chuxiong, Yunnan province, China, susceptibility to buprofezin increased after removal of Wolbachia. An increase in susceptibility was also evident in a Wolbachia-infected line established from a field population in Rugao, Jiangsu province. However, no increase was evident in two field populations from Nanjing and Fengxian, Jiangsu province, China. When Wolbachia was introgressed into different genetic backgrounds, followed by Wolbachia removal, the data pointed to Wolbachia effects that depend on the nuclear background as well as on the Wolbachia strain. However, there was no relationship between Wolbachia density and the component of buprofezin resistance associated with This article is protected by copyright. All rights reserved. 4 the symbiont. The results suggest that Wolbachia effects associated with chemical resistance are complex and unpredictable, but also that they can be substantial.
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