Using stable isotope analysis to examine fall armyworm (Lepidoptera: Noctuidae) host strains in a cotton habitat

Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), or fall armyworm, is an important agricultural pest of several crops in the Western Hemisphere, including cotton (Gossypium L.). Two morphologically identical host strains of fall armyworm exist that differ in plant host use and habitat distribution. The corn-strain is a primary pest of corn, Zea mays L., whereas the rice-strain is the majority population infesting rice (Oryza spp.) and turfgrass (Cynodon spp.). With the increased use of Bacillus thuringiensis (Bt) toxin-expressing cotton varieties and the necessity of ensuring adequate refuge areas to prevent the spread of Bt toxin resistance, it is crucial to identify the alternative plant hosts available for the fall armyworm population infesting cotton. Stable isotope analysis combined with the molecular analysis of strain-specific markers was used to investigate whether one or both strains routinely develop on cotton grown in the Mississippi delta. We found that the majority of fall armyworm adults present during the early cotton growing season arose from C4 plants (e.g., corn and sorghum, Sorghum vulgare Pers.) and that the only strain likely to be developing on cotton (a C3 plant) in substantial numbers was the corn-strain. The population distribution patterns observed were consistent with corn providing an important refuge for the fall armyworm strain infesting cotton and suggested that late season populations in the Mississippi delta may be migrants from more northern corn areas.     

Genetic characterization of fall armyworm (Lepidoptera: Noctuidae) host strains in Argentina

Fall armyworm is a major economic pest throughout the Western Hemisphere. Previous studies of populations in the southern United States, Brazil, and the Caribbean demonstrated the existence of two morphologically identical but genetically distinct host strains that can only be distinguished using genetic markers, including polymorphisms in the mitochondrial Cytochrome Oxidase I (COI) gene and in the Z-chromosome linked Triose phosphate isomerase (Tpi) gene. The strains differ in some physiological and behavioral characteristics, most notably their preference for different plant hosts, but are capable of hybridizing in the laboratory and in the field. These traits suggest that the strains are in the process of divergence, which may or may not be hemispheric in scope. The objective of this study was to determine whether the two strains are present in Argentina. It was found that the strain-diagnostic haplotypes of the COI and Tpi genes subdivided the Argentina population into two major groups. Each group displayed biases in their distribution among different host plants that were generally consistent with expected strain behavior. The overall results indicated that Argentina fall armyworm exhibit similar genetics and behavior to populations in the rest of the hemisphere. In addition, the Argentina populations had comparable haplotype frequencies to those from Brazil and Texas, consistent with possible interactions with these fall armyworm groups, but appeared to have had minimal exchanges with those from Puerto Rico or Florida.     

New restriction fragment length polymorphisms in the cytochrome oxidase I gene facilitate host strain identification of fall armyworm (Lepidoptera: Noctuidae) populations in the southeastern United States

Several restriction sites in the cytochrome oxidase I gene of fall armyworm, Spodoptera frugiperda (J.E. Smith), were identified by sequence analysis as potentially being specific to one of the two host strains. Strain specificity was demonstrated for populations in Florida, Texas, Mississippi, Georgia, and North Carolina, with an AciI and SacI site specific to the rice (Oryjza spp.)-strain and a BsmI and HinfI site joining an already characterized MspI site as diagnostic of the corn (Zea mays L.)-strain. All four of these sites can be detected by digestion of a single 568-bp polymerase chain reaction-amplified fragment, but the use of two enzymes in separate digests was found to provide accurate and rapid determination of strain identity. The effectiveness of this method was demonstrated by the analysis of almost 200 adult and larval specimens from the Mississippi delta region. The results indicated that the corn-strain is likely to be the primary strain infesting cotton (Gossypium spp.) and that an unexpected outbreak of fall armyworm on the ornamental tree Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud. was due almost entirely to the rice-strain.     

基于分子标记的入侵陕西草地贪夜蛾种群生物型分析

草地贪夜蛾Spodoptera frugiperda J. E. Smith是一种世界性入侵害虫,为了明确入侵陕西的草地贪夜蛾的种群生物型并了解其发生及扩散规律,本研究对采集自陕西省8个地市180个草地贪夜蛾样本分别进行了基于COI和Tpi分子标记的生物型鉴定。分析发现入侵陕西的草地贪夜蛾84%是水稻型母本与玉米型父本杂交形成的杂合玉米型草地贪夜蛾。COI分子标记的结果显示,样本中水稻型占比为84.44%,玉米型为15.56%;基于Tpi基因片段的结果表明除商洛样本SL-3外,其它样本均为玉米型。值得注意的是,SL-3与非洲特异型序列同源性达100%,此非洲特异单倍型为陕西省内首次、国内第2次出现。本研究为草地贪夜蛾的迁飞扩散规律及早期预警提供了新的理论基础。     

Using haplotypes to monitor the migration of fall armyworm (Lepidoptera: Noctuidae) corn-strain populations from Texas and Florida

Fall armyworm, Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae), infestations in most of North America north of Mexico arise from annual migrations of populations that overwinter in southern Texas and Florida. A comparison of the cytochrome oxidase I haplotype profiles within the fall armyworm corn-strain, the subgroup that preferentially infests corn (Zea mays L.) and sorghum (Sorghum vulgare Pers.), identified significant differences in the proportions of certain haplotypes between the Texas and Florida populations. These proportional differences were preserved as the populations migrated, providing a molecular metric by which the source of a migrant population could be identified. The migratory pattern derived from this method for several southeastern states was shown to be consistent with predictions based on analysis of historical agricultural and fall armyworm infestation data. These results demonstrate the utility of haplotype proportions to monitor fall armyworm migration, and they also introduce a potential method to predict the severity of cotton crop infestations in the short term.     

Influence of plant size and species on preference of Cyrtobagous salviniae adults from two populations

Adults from two populations (Brazil and Florida) of Cyrtobagous salviniae were bioassayed to determine if they exhibited a preference for either Salvinia minima or Salvinia molesta. Adults did not discriminate between host species in initial tests that evaluated the tertiary growth form. Further tests which compared two growth forms (primary and tertiary) as well as plant species, found that adults from the Brazil population consistently preferred larger (tertiary) plants without regard for host species. Weevils from the Florida population showed a similar, but less distinct, pattern of preference. Although adults from the Florida population survived equally well and experienced a similar pre-oviposition period on both plant species, they laid more eggs in S. molesta. Adults from the two populations differ in size: Brazil weevils were larger, which may explain their sensitivity to plant size as compared with the smaller Florida adults. Narrower rhizomes in S. minima may restrict usage of this species by the larger weevils, whereas smaller larvae may be better able to burrow in a wider range of plant sizes. Both weevil populations should be suitable biological control agents for use in programs targeting S. molesta.     

Susceptibility of Spodoptera frugiperda (Lepidoptera: Noctuidae) strains from central Colombia to two insecticides, methomyl and lambda-cyhalothrin: a study of the genetic basis of resistance

Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae) is an important pest of several crops in the western hemisphere. This insect has genetically differentiated into two host-associated populations: the corn (Zea mays L.) and the rice (Oryza sativa L.) strains. The corn strain also is found in cotton (Gossypium hirsutum L.) and sorghum and the rice strain in Bermuda grass [Cynodon dactylon (L.) Pers.] and millet. In the United States and Brazil, lines from corn, rice, Bermuda grass, and millet were used to evaluate the resistance of both strains to various insecticides, and found that the corn strain is more resistant than the rice strain. However, in these studies the larvae were not genotyped. In Colombia, genotyping of fall armyworm is necessary because the rice strain also can be found in corn fields. In this work, collected larvae from corn and rice fields from Tolima (central Colombia) were genotyped and evaluated for the resistance to methomyl and lambda-cyhalothrin. We found that the rice strain does not significantly differ in resistance to methomyl compared with the corn strain but it develops tolerance more rapidly to lambda-cyhalothrin. The eggs viability of treated females also was significantly affected by methomyl on each generation. The realized heritability of resistance was higher for lambda-cyhalothrin (0.23-0.42) than for methomyl (0.04-0.14). The number of generations needed for 10-fold increase in resistance is approximately 11.5 generations for methomyl and 6.5 for lambda-cyhalothrin. Finally, the genetic basis of resistance to both insecticides involves few recessive autosomal genes. The results obtained here suggest that methomyl is a better option than lambda-cyalothin to control fall armyworm.     

Effects of defoliation on the resistance and tolerance of rice,Oryza sativa,to root injury by the rice water weevil,Lissorhoptrus oryzophilus

Understanding plant‐mediated interactions in agricultural systems may facilitate the development of novel and improved management practices, which is important, as management of these insects is currently heavily reliant on insecticides. The fall armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae, Prodeniini), is a sporadic pest of rice fields in the southern USA. In southwestern Louisiana, this defoliating insect typically attacks rice early in the growth season, before fields are flooded. Defoliation by fall armyworm larvae may trigger increased expression of plant defenses, which may result in increased resistance to subsequent herbivores. The rice water weevil (RWW), Lissorhoptrus oryzophilus Kuschel (Coleoptera: Curculionidae, Stenopelmini), enters rice fields as an adult both before and after flooding, but oviposition and larval infestation occur only after fields are flooded. RWW may be affected by changes in plant resistance caused by fall armyworm defoliation before flooding. The objectives of this study were to investigate the plant‐mediated effects of natural and artificial defoliation on population densities of RWW larvae after flooding and on the ability of rice plants to compensate for root injury by RWW larvae. In the 2015 season, fall armyworm defoliation before flooding resulted in reduced RWW densities after flooding. However, in 2016 no significant effects of fall armyworm defoliation on densities of RWW larvae were detected. Similarly, mechanical defoliation of rice before flooding did not affect RWW densities after flooding. Although lowest yields were observed in plots subjected to both root injury and defoliation, there was little evidence of a greater than additive reduction in yields from simultaneous injury. These results suggest a lack of plant‐mediated interactions among these two pests in rice in the southern USA.     

Factoring distribution and prevalence of Fall armyworm in southwest China

Host plant growth changing with environmental conditions can impact the distribution of herbivores. The generalist herbivore fall armyworm (FAW), Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae), is an invasive pest rapidly spreading across the world and has recently invaded southern China. We studied effects of environmental factors on the distribution of the fall armyworm and its host (maize) plant growth in the tropical mountainous area of Huize County, province of Yunnan, southwest China. Moreover, the relationships among the FAW distribution, environmental factors (altitude, temperature and humidity) and plant growth (mean kernel weight, kernel number per ear and ear weight) were analysed. The results showed that FAW predominated at altitude 1,243.3 m, temperature 21.4°C and humidity 82.1%. The host plant grew best at 1,200–1,312 m, 21.0–21.7°C and 81.2%–82.0%. Environmental factors indirectly influenced the distribution of FAW via the host plant growth. Compared with environmental factors, the host plant growth had a simplistic positive linear relationship with the density of FAW. FAW is less impacted by abiotic factors rather it determined by host plant prevalence, and thus the locations where maize plants grow best are preferred by FAW and should be the focus of insecticide applications. Understanding the distribution of FAW under various environmental conditions provides a valuable reference for Chinese maize production and food security.     

Subtropical Interactions: Comparing Galling Insect and Host Plant Diversity in Southern Brazil and Florida

Gall-inducing insects seem to have a diversity pattern distinct from the usual latitudinal decrease in species, with more species occurring in xeric environments instead. Many questions regarding galler diversity over geographical scales remain unanswered: for example, little is known about beta diversity, and the role super host plants play in local/regional richness. Our aim was to compare galling insect and host plant diversity in different biogeographical regions, but under similar environmental conditions. We sampled short stature coastal woodlands on sandy soils of the Atlantic coast in both USA (Florida) and Brazil (Rio Grande do Sul, RS), between 25° and 30° latitude. Little-used 200-m long trails were searched during 90 min for galls; there were four trails in USA and five in Brazil. Gall functional traits (galled plant organ, gall shape and colour) proportions were not different between Florida and RS. Local galling and host plant species richness also did not differ, and neither did regional galling diversity. The beta diversity pattern, however, was distinct: sites in Florida have more similar galling faunas than sites in RS. Common diversity patterns indicate common environmental biotic (plant diversity, vegetation structure) and abiotic (climate, soil) factors might be contributing to these similar responses. As Brazilian sites are in the Atlantic forest hotspot, a high galling insect beta diversity might be caused by a higher heterogeneity at larger scales—sample-based rarefaction curves were ascending for Brazil, but not for USA. Myrtaceans were super hosts in Brazil, but not in Florida, where oaks take up this role.     

不同生育期玉米上草地贪夜蛾的发生为害规律

2019年在广东湛江、江门调查了玉米上草地贪夜蛾Spodoptera frugiperda (Smith)发生为害动态,分析了不同生育期玉米上该虫发生为害差异规律。发现各个地区草地贪夜蛾的虫口数量与玉米受害程度存在较大差异,呈现不均衡性;该虫对玉米苗期为害最严重,其次为大喇叭口期;建立了2个地区玉米6个生育期受害率与虫口数量之间的线性回归模型。研究结果可为草地贪夜蛾监测预警及防控提供参考。     

Population genetic structure of two primary parasitoids of Spodoptera frugiperda (Lepidoptera), Chelonus insularis and Campoletis sonorensis (Hymenoptera): to what extent is the host plant important?

Plant chemistry can strongly influence interactions between herbivores and their natural enemies, either by providing volatile compounds that serve as foraging cues for parasitoids or predators, or by affecting the quality of herbivores as hosts or prey. Through these effects plants may influence parasitoid population genetic structure. We tested for a possible specialization on specific crop plants in Chelonus insularis and Campoletis sonorensis, two primary parasitoids of the fall armyworm, Spodoptera frugiperda. Throughout Mexico, S. frugiperda larvae were collected from their main host plants, maize and sorghum and parasitoids that emerged from the larvae were used for subsequent comparison by molecular analysis. Genetic variation at eight and 11 microsatellites were respectively assayed for C. insularis and C. sonorensis to examine isolation by distance, host plant and regional effects. Kinship analyses were also performed to assess female migration among host‐plants. The analyses showed considerable within population variation and revealed a significant regional effect. No effect of host plant on population structure of either of the two parasitoid species was found. Isolation by distance was observed at the individual level, but not at the population level. Kinship analyses revealed significantly more genetically related—or kin—individuals on the same plant species than on different plant species, suggesting that locally, mothers preferentially stay on the same plant species. Although the standard population genetics parameters showed no effect of plant species on population structure, the kinship analyses revealed that mothers exhibit plant species fidelity, which may speed up divergence if adaptation were to occur.     

Quantitative proteomic analysis of the fall armyworm saliva

Lepidopteran larvae secrete saliva on plant tissues during feeding. Components in the saliva may aid in food digestion, whereas other components are recognized by plants as cues to elicit defense responses. Despite the ecological and economical importance of these plant-feeding insects, knowledge of their saliva composition is limited to a few species. In this study, we identified the salivary proteins of larvae of the fall armyworm (FAW), Spodoptera frugiperda; determined qualitative and quantitative differences in the salivary proteome of the two host races—corn and rice strains—of this insect; and identified changes in total protein concentration and relative protein abundance in the saliva of FAW larvae associated with different host plants. Quantitative proteomic analyses were performed using labeling with isobaric tags for relative and absolute quantification followed by liquid chromatography-tandem mass spectrometry. In total, 98 proteins were identified (>99% confidence) in the FAW saliva. These proteins were further categorized into five functional groups: proteins potentially involved in (1) plant defense regulation, (2) herbivore offense, (3) insect immunity, (4) detoxification, (5) digestion, and (6) other functions. Moreover, there were differences in the salivary proteome between the FAW strains that were identified by label-free proteomic analyses. Thirteen differentially identified proteins were present in each strain. There were also differences in the relative abundance of eleven salivary proteins between the two FAW host strains as well as differences within each strain associated with different diets. The total salivary protein concentration was also different for the two strains reared on different host plants. Based on these results, we conclude that the FAW saliva contains a complex mixture of proteins involved in different functions that are specific for each strain and its composition can change plastically in response to diet type.     

Parallel host shifts in a bacterial plant pathogen suggest independent genetic solutions

While there are documented host shifts in many bacterial plant pathogens, the genetic foundation of host shifts is largely unknown. Xylella fastidiosa is a bacterial pathogen found in over 600 host plant species. Two parallel host shifts occurred—in Brazil and Italy—in which X. fastidiosa adapted to infect olive trees, whereas related strains infected coffee. Using 10 novel whole-genome sequences from an olive-infecting population in Brazil, we investigated whether these olive-infecting strains diverged from closely related coffee-infecting strains. Several single-nucleotide polymorphisms, many derived from recombination events, and gene gain and loss events separated olive-infecting strains from coffee-infecting strains in this clade. The olive-specific variation suggests that this event was a host jump with genetic isolation between coffee- and olive-infecting X. fastidiosa populations. Next, we investigated the hypothesis of genetic convergence in the host shift from coffee to olive in both populations (Brazil and Italy). Each clade had multiple mutations and gene gain and loss events unique to olive, yet no overlap between clades. Using a genome-wide association study technique, we did not find any plausible candidates for convergence. Overall, this work suggests that the two populations adapted to infect olive trees through independent genetic solutions.     

Spatial distribution of plants infested with Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) on corn crop

The importance of the fall armyworm, Spodoptera frugiperda (J.E. Smith), is not only due to the damage it can cause, but also to its difficult control. It is essential to know the pest population parameters, such as its standard scattering in the crop, because depending on the space arrangement of the insects in the area, different sampling methods are required. For the present work, three experimental fields were installed in different areas: the first one had 66 plots with seven plants each; the second one had 55 plots with four plants each; and the third field had 55 plots with six plants each. The presence or absence of the pest in the plant was evaluated in the samples. According to the data observed, either to the scattering index or to the theoretical scattering models, S. frugiperda dispersion is scattered at random and strongly depends on the infestation level.     

Cover Caption

The fall armyworm, Spodoptera frugiperda, is a major pest native to the Americas that has recently invaded the Old World. A whole-genome sequencing (WGS) approach was used to identify mutations associated with resistance to insecticides and Bt proteins in 150 individuals of S. frugiperda collected from China, Malawi, Uganda and Brazil. This approach revealed three amino acid substitutions (A201S, G227A and F290V) of the acetylcholinesterase-1 (ace-1) in the four geographical populations of S. frugiperda, which are known to be associated with organophosphate resistance. A novel Cry1F resistance allele in the ABCC2 gene was identified from the Brazilian population but absent in the invasive populations (see pages 627-638). Photo provided by Prof. Run-Zhi Zhang.     

Trajectory modeling revealed a southwest-northeast migration corridor for fall armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) emerging from the North China Plain

The fall armyworm Spodoptera frugiperda, an invasive insect pest native to the Americas, has established populations throughout eastern China. The North China Plain—a key corn-producing area in East China with a unique topography—was invaded by fall armyworm in 2019 and is seriously threatened by this migratory pest. However, the spatiotemporal extent of the migratory movements of fall armyworm from the North China Plain remains poorly understood. Using an air transport-based trajectory modeling approach that incorporates flight behavior, we simulated the potential nocturnal migration trajectories of fall armyworm from the North China Plain based on historical meteorological data from June to October of 2015–2019, and examined the night-time atmospheric conditions associated with their possible flights. The emigration patterns showed monthly variation in the main landing area and common migration direction. The displacement of newly emerged moths from the North China Plain was concentrated in the Northeast China Plain (including Liaoning, Jilin and Heilongjiang provinces) before late summer, after which they were most likely to undertake return flights to the south (especially into Hubei, Anhui and Hunan provinces). This southwest-northeast aerial migration corridor follows the topography of East China and is affected by the East Asian monsoon. These topographic-atmospheric conditions have resulted in the North China Plain becoming a key stopover for fall armyworm populations engaging in multigenerational long-distance migration across East China. These findings contribute to our knowledge of fall armyworm migration and will aid in the implementation of management and control strategies against this highly migratory agricultural pest.     

MtDNA variability in whitefly (Bemisia tabaci) populations in Brazil

Bemisia tabaci (Hemiptera: Aleyrodidae) consists of a complex of morphologically indistinct biotypes that vary mainly in their capacity to transmit plant viruses and to induce physiological disorders in plants of economic importance. The adaptability of B. tabaci to many regions of the world has fostered the appearance of various biotypes and has resulted in a broad spectrum of host plants. Our goal was to identify which biotypes were present in four B. tabaci populations in Brazil. We quantified genetic variability between and within populations. Three individuals were collected from three host plant species: two populations on soybean (Campinas and Rondonópolis), one on pumpkin (Barreiras) and one on tomato (Cruz das Almas) in three States of Brazil (S?o Paulo, Mato Grosso, and Bahia). We chose one sequence of the B biotype, obtained from GenBank; the Campinas population, which had been previously characterized as biotype B, was used as a control for this biotype. We also included one sequence of the Q biotype, obtained from GenBank, as an outgroup. The COI region of the mtDNA gene was partially amplified with the CI-J-2195 and L2-N-3014 pair of primers, and the reaction products were sequenced. Based on distance-based algorithm analyses, we found that all haplotypes belong to biotype B, which was confirmed by the haplotype network. Genetic structure analyses showed that the host plant species does not influence population structuring of this pest; only the geographic location mattered.