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.     

Oviposition Choice of Two Fall Armyworm (Lepidoptera: Noctuidae) Host Strains

Fall armyworm, Spodoptera frugiperda (J. E. Smith), is a noctuid species that is composed of two morphologically identical sympatric host strains (corn and rice) that differ in their distribution among plant hosts. In an effort to explain observations of host fidelity in the field, ovipositional preference of the two strains on corn (Zea mays L.) or pasture grass (Cynodon nlemfuensis Vanderyst var. nlemfuensis) was determined using two greenhouse bioassays. In the first bioassay, corn strain females placed more eggmasses on the screen enclosure than on corn plants while grass plants contained an intermediate number of eggmasses. Rice strain females placed most of their eggmasses on grass plants. In the second bioassay, corn strain females placed an equal number of eggmasses on corn and grass plants in comparison to rice strain females which placed >3.5× more eggmasses on grass plants than on corn plants. Individual eggs as part of the eggmasses were also counted on plants and on the screen enclosure. Corn strain females equally placed eggs on the two host plants and on the screen enclosures, however rice strain females placed more eggs on grass plants compared to corn plants or the screen enclosure. This is the first report of consistent differential oviposition between corn and rice strain fall armyworm females.     

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.     

QUANTITATIVE GENETICS,DEVELOPMENT, AND PHYSIOLOGICAL ADAPTATION IN HOST STRAINS OF FALL ARMYWORM

Two genetically differentiated host strains of fall armyworm were reared on their own and each other's host plants, rice and corn, to determine whether they were physiologically adapted to their natural hosts and whether they exhibited genetically based differences in development. Larval host had a greater impact on development in the rice strain than in the corn strain, indicating that physiology could have facilitated specialization in one strain but not the other. Consequently, behavioral mechanisms are also likely to be important in the maintenance of host specificity. Comparisons between strains indicated significant differences in one trait, the rate at which larvae gained weight. Because this character had consistently high heritabilities, genetic differentiation in development is indicated. An analysis of genotype-by-environment interactions within each strain detected significant interactions for three of five traits, suggesting that genotypic performance on one host was not indicative of performance on the other. Each strain thus exhibited genetic variation that would facilitate host-associated divergence and adaptation if coupled with a mechanism that reduced gene flow between hosts. Finally, significant genetic correlations between several characters were detected when strains were reared on their natural hosts but not when they were reared on nonnatural hosts. Apparently, feeding on novel hosts caused developmental uncoupling of characters. Release from genetic constraints could provide a mechanism for physiological adjustments to newly occupied habitats.     

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.     

Comparison of pheromone trap design and lures for Spodoptera frugiperda in Togo and genetic characterization of moths caught

Fall armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae), is a pest of grain and vegetable crops endemic to the Western Hemisphere that has recently become widespread in sub‐Saharan Africa and has appeared in India. An important tool for monitoring S. frugiperda in the USA is pheromone trapping, which would be of value for use with African populations. Field experiments were conducted in Togo (West Africa) to compare capture of male fall armyworm using three commercially available pheromone lures and three trap designs. The objectives were to identify optimum trap × lure combinations with respect to sensitivity, specificity, and cost. Almost 400 moths were captured during the experiment. Differences were found in the number of S. frugiperda moths captured in the various trap designs and with the three pheromone lures, and in the number of non‐target moths captured with each lure. The merits of each trap × lure combination are discussed with respect to use in Africa. A nearly equal number of COI‐CS (161) and COI‐RS (158) moths was captured with no differences found in COI marker proportions among traps or lures. However, the diagnostic rice strain marker Tpi was rarely found. Overall, the genetic characterization of the pheromone trap collections indicated a consistent distribution of genetic markers from 2016 to 2017, suggesting a population at or near equilibrium.     

Attractiveness of binary blends of floral odorant compounds to moths in Florida, USA

Evaluation of combinations of flower odor compounds in the field revealed several compounds that were attractive or co-attractive with phenylacetaldehyde (PAA) to pest noctuid and pyralid moths. A number of moth species responded positively to the key floral odorant PAA. The floral odorants cis -jasmone, linalool, benzyl acetate, limonene, β-myrcene, methyl salicylate, and methyl 2-methoxybenzoate all increased captures of some moths when added to traps with PAA, but responses varied among the moth species that were trapped. For example, soybean looper moths, Pseudoplusia includens (Walker) (Lepidoptera: Noctuidae), most strongly responded to PAA + β-myrcene, but benzyl acetate, cis -jasmone, and limonene also increased captures of these moths when these compounds were used in traps along with PAA. Velvetbean caterpillar moths, Anticarsia gemmatalis Hübner (Lepidoptera: Noctuidae), responded most strongly to PAA + linalool, but β-myrcene, cis -jasmone, and limonene also increased captures of these moths in traps over numbers trapped with PAA. Positive responses to floral compound blends were also noted for golden looper [ Argyrogramma verruca (F.)], grass looper [ Mocis disseverans (Walker)], tobacco budworm [ Heliothis virescens (F.)], southern armyworm [ Spodoptera eridania (Stoll) (all Lepidoptera: Noctuidae)], and melonworm [ Diaphania hyalinata (L.) (Lepidoptera: Pyralidae)]. Overall, PAA + β-myrcene appeared to be the strongest floral compound combination for pest species trapped, compared to PAA or other compound blends.     

Downwind migration of the African army worm moth, Spodoptera exempta, studied by mark-and-capture and by radar

Abstract. 1. About 166,000 African armyworm moths, Spodoptera exempta (Walk.), were marked at an emergence site near Nairobi when they fed at night on trees baited with dyed molasses.
2. Six marked moths were captured in pheromone traps, including one at 90 km after flying for only one night, and another at 147 km.
3. Moth flight trajectories deduced from radar and from marking showed that migration was downwind.
4. During migration, moths become dispersed; hence the high densities that lead to outbreaks must be produced by concentration.
5. Some moths were ready to mate on the same night they completed their long-distance flight.     

Changes in distribution of an archaic moth, Micropterix calthella, in St Petersburg, Russia, between 1989 and 2005

Aim  This work aimed to assess changes in the number of suitable habitats and habitat occupancy by Micropterix calthella (L.) moths (Lepidoptera, Micropterigidae) in the city of St Petersburg between 1989 and 2005.
Location  The city of St Petersburg, Russia.
Methods  The study compares results of the survey conducted on 9–12 June 2005 with data collected in 1989 and is based on the assessment of the quality and occupancy of 103 habitats that were found potentially suitable for M. calthella in 1989. If habitats contained wet microsites with undisturbed litter and dense field layer vegetation with a high abundance of flowering Ranunculus sp., they were considered potentially suitable for M. calthella ; in these habitats, moths were searched for on flowers of Ranunculus – visually and/or by extensive sweep-netting.
Results  Both the number of habitats suitable for M. calthella and the number of habitats occupied by this species decreased since 1986 from 103 to 72 and from 58 to 34, respectively. The disappearance of habitats was primarily due to the building of residential blocks within the city (16 sites) and railway renovation (10 sites), while changes in habitat management, primarily in parks and recreation zones (five sites), were less important. The average distance from the city centre to the nearest population of M. calthella increased from 8.7 ± 1.5 to 12.9 ± 3.0 km (mean ± SE).
Main conclusion  The data for 2005 fit the prediction that M. calthella moths, which are unable to re-colonize potentially suitable habitats due to their low migratory ability, will disappear from the city. At the present rate of extinction, this process may take some 20–30 years to complete (or nearly complete).     

Comparison of haplotype frequencies differentiate fall armyworm (Lepidoptera: Noctuidae) corn-strain populations from Florida and Brazil

Fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), is a major economic pest throughout the Western Hemisphere. Populations can be subdivided into two morphologically identical but genetically distinct strains (corn-strain and rice-strain) that differ in their host plant preferences. These strains can be distinguished by using polymorphisms in the mitochondrial cytochrome oxidase 1 gene. Additional sequence analysis of this locus identified two sites that were highly polymorphic in the corn-strain population and that produced four different haplotype subgroups. Comparisons of the frequency distribution of these haplotypes found no seasonal or plant host specificities, but they did demonstrate that the Brazil corn-strain population is different from corn-strain fall armyworm found in Florida. The development of a rapid means of distinguishing fall armyworm populations originating from Brazil versus Florida provides an opportunity for investigating and comparing the genetic complexity and long-range movements of this important agricultural pest.     

Migration and Allee effects in the six-spot burnet moth Zygaena filipendulae

Abstract 1. Migration into local populations may increase the likelihood of persistence but emigration may decrease the persistence of small and isolated populations. The dispersal behaviour of a day-flying moth Zygaena filipendulae was examined to determine whether emigration is correlated positively or negatively with population size and host plant density.
2. A mark–release–recapture study showed that most moths moved small distances (< 40 m on average) and only 6% of movements were > 100 m.
3. Twenty-five individuals moved between populations, a measured exchange rate of 8%. Moths were more likely to move between patches that were close together and they moved to relatively large patches.
4. The fraction of residents increased with increasing population size in the patch and increasing host plant cover. Relatively high proportions of individuals left small patches with small moth populations.
5. Moths released in grassland lacking Lotus corniculatus (the host plant) tended to leave the area and biased their movement towards host plant areas, whereas those released within an area containing L. corniculatus tended to stay in that area.
6. Biased movement away from small populations and areas of low host plant density (normally with low population density) was found. This migration-mediated Allee effect is likely to decrease patch occupancy in metapopulations, the opposite of the rescue effect. The effects on metapopulation persistence are not known.     

Time‐shifted reproductive behaviours among fall armyworm (Noctuidae: Spodoptera frugiperda) host strains: evidence for differing modes of inheritance

The noctuid moth Spodoptera frugiperda consists of two strains associated with different larval host plants (most notably corn and rice). These strains exhibit differential temporal patterns of female calling and copulation during scotophase, with the corn strain more active earlier in the night. We investigated strain‐specific constraints in reproductive timing, mating interactions between the two strains, and the mode of inheritance of timing of female calling, male calling, copulation and oviposition. We observed an allochronic shift of all reproductive behaviours by approximately 3 h and a parallel shift of nonreproductive locomotor activity, suggesting involvement of the circadian clock. The corn strain was more variable in the timing of calling and copulation than the rice strain. Rice strain females were more restricted in the timing of copulation than rice strain males, while such differences between the sexes were not apparent in the corn strain. There were significant interactions between the strains affecting onset times of copulation and male calling. The four investigated reproductive traits differed in their modes of inheritance: timing of female and male calling exhibited strong maternal effects, timing of copulation was controlled by a combination of maternal effects and corn strain dominant autosomal factors, and timing of oviposition was inherited in a corn strain dominant fashion. We conclude that the allochronic separation of reproduction between fall armyworm strains is asymmetric, less pronounced than previously thought, and under complex genetic control.     

广东省草地贪夜蛾冬季发生特征及周年繁殖区域研究

广东省是草地贪夜蛾入侵我国的桥头堡和主要的北迁虫源地之一,明确该虫在广东省的冬季发生特征及越冬存活情况,对广东省乃至全国草地贪夜蛾的预测预报及源头治理意义重大。为准确掌握广东省草地贪夜蛾的周年繁殖区范围及冬季发生为害情况,2020年1-3月在粤东、粤西、粤北及珠三角地区,利用成虫性诱、挖土查蛹、幼虫密度及植株为害率普查等方法,分析广东省草地贪夜蛾冬前、冬后种群发生为害情况及冬季发生特征。调查结果表明:(1)广东省冬玉米种植区主要分布在湛江、茂名、阳江、惠州等地,冬玉米种植区均发现草地贪夜蛾幼虫为害,主要为害冬玉米,极少为害甘蔗;(2)不同地区冬种玉米上草地贪夜蛾的发生程度差异较大,湛江、茂名、阳江发生为害较为严重,平均为害率30%左右,而珠三角及粤东地区发生较轻,为害率低于10%;较冬前调查,冬后草地贪夜蛾发生量和为害程度出现不同程度的下降;(3)广东省大部分地区的冬种玉米田和空闲地均可持续诱捕到草地贪夜蛾成虫,而挖土调查发现草地贪夜蛾蛹密度较低。本调查明确了草地贪夜蛾在北回归线以南的冬玉米区可以周年繁殖,无明显的滞育越冬现象,粤西茂名、阳江以南至雷州半岛一带为典型冬种玉米区,草地贪夜蛾发生为害较为严重,珠三角及粤东大部分地区草地贪夜蛾种群数量相对较低,调查结果为广东省乃至全国草地贪夜蛾的早期预警和精准防控提供了重要参考。     

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.     

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.     

Quantification of diapausing fourth generation and suicidal fifth generation cotton bollworm, Helicoverpa armigera, in cotton and corn in northern China

Cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), is a major pest of cotton and corn crops in northern China. A phenological differentiation between fourth generation cotton bollworms from cotton and those from corn fields was observed in northern China during 1999–2000. The proportion of pupation in late fall was marginally higher in cotton fields compared to that in corn fields; however, the proportions of fall emergence of moths from cotton fields were significantly higher than those from corn fields. The proportion of spring emergence of moths was also significantly higher for larvae collected from cotton (28.0%) than from corn (14.5%). The overwintering duration of females was significantly shorter than that of males in both crops. Moreover, the overwintering duration of bollworm populations from cotton was significantly longer than that from corn. The early spring population of H. armigera came from both cotton and corn fields, but the spring emergence of moths from larvae collected from cotton took about 5 days longer to reach 100% emergence compared to that from corn.     

Host association of Spodoptera frugiperda (Lepidoptera: Noctuidae) corn and rice strains in Argentina, Brazil, and Paraguay

Spodoptera frugiperda (J.E. Smith) is composed of two genetically distinct strains, the so-called corn strain and the rice strain. Whether the two strains differ in their host use is unclear, because laboratory experiments have not been able to show consistent host performance or preference differences between them, and field studies showed high rates of hybridization, as well as some degree asymmetric host use. To determine the distribution of the two strains and their association with host plants, we collected fall armyworm larvae from different crops (corn, rice, alfalfa, and sorghum) and grasses in 15 different localities over 4 yr in Argentina, Brazil, and Paraguay. The strain identity was analyzed using two polymorphisms in the mitochondrial cytochrome oxidase subunit I gene. We identified the corn and rice haplotypes and three types of populations were characterized based on the frequencies of the individuals that belonged to any of these haplotypes: in 44% of populations the corn haplotype predominated, in 44% of populations the rice haplotype was the most frequent, and 11% of populations showed both haplotypes at similar proportions. In total, eight populations (47%) showed the expected pattern, two populations (12%) were polymorphic within the same field, and seven populations (41%) showed the inverse pattern. Taken together, there was no consistent pattern of host association between the two sympatric genotypes and their respective host plants. This investigation supports the need for additional studies to determine which other forces keep the genotypes separate, and what is the degree of genetic differentiation between these populations.     

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 intergenerational host plant alterations on developmental parameters of Spodoptera frugiperda (Lepidoptera: Noctuidae)

The fall armyworm (FAW), Spodoptera frugiperda (Smith), is known as a destructive moth pest of agricultural crops such as corn, rice, and sorghum. Recently, its damage threatens crops in Asian temperate regions due to its high migration ability. Two genetic types of FAW, a corn strain mostly found on corn and a rice strain associated with rice, have been reported. The objective of this study was to evaluate whether migrated corn-collected population of FAW could maintain its population on various seasonal plants including non-preferred Poaceae plants, with special emphasis on Jeju, Korea. Development-related parameters were compared after switching host plants including corn (C), Sudan grass (S), proso millet (M), and rye (R) when rearing FAW. Most developmental parameters showed dynamic changes according to sequential alternation of host plants with the progress of generations. When all R₀ (net reproductive rate) values were scaled by the R₀ in C to C alteration, the scaled values of other alterations were less than “1.0”, indicating less fitness than C to C alteration (R₀ value: 0.57 for C to S, 0.49 for C to M in F3 generation; 0.82 for S to S, 0.65 for S to M, and 0.58 for S to C in F4 generation, 0.50 for M to M, 0.59 for M to R, and 0.83 for M to C in F5 generation). Although our results did not show that S. frugiperda adapted perfectively on new host plants, this pest was able to successfully reproduce up to five generations with non-preferred host plants. Results indicate that winter crops such as rye might be hosts for migrated corn-preferred populations of S. frugiperda during winter after breeding three to four cycles on other host plants before winter if weather condition is allowed for its survival.