Migration of invasive Spodoptera frugiperda(Lepidoptera: Noctuidae) across the Bohai Sea in northern China

The fall armyworm(FAW), Spodoptera frugiperda(J. E. Smith), is an important agricultural pest with a strong migratory ability. While the species is native to the Americas, it has recently invaded China, ravaging crops in many provinces(autonomous regions, municipalities). Its long-distance migration, which is critical to understand for pest management programs, has been well studied in its native region. In other regions however, the moth's migration patterns have not yet been characterized. Here, the migratory behavior of FAW populations in China was studied on an isolated island located in the center of the Bohai Strait from spring to autumn 2019, the year in which FAW first reached this region, by using searchlight trapping, stable carbon isotopes and trajectory simulation. The main results were summarized as follows:(i) The number of FAW moths caught by searchlight trapping provided direct evidence that the species migrated across the Bohai Sea.(ii) Species identification was confirmed by both morphology and molecular methods, and only the "corn-strain" that preferentially infests maize and sorghum was found in the collections.(iii) Stable carbon isotope measurements showed that up to 95.07% of captured moths displayed the C4 isotope signature, thus providing additional evidence that this species is a migrant as there are no major C4 plants at the trapping site.(iv) Backward trajectory analysis indicated that the pest now threatens to expand its range into the agriculturally important region of Northeast China, and thus authorities in this region should be vigilant to the threat posed by this species. Together, these findings add to our knowledge concerning the occurrence of FAW in northern China, and will help us to develop sustainable and effective monitoring, forecasting, and pest management strategies.     

Windborne migration routes of newly-emerged fall armyworm from Qinling Mountains–Huaihe River region,China

The fall armyworm(FAW, Lepidoptera: Noctuidae), Spodoptera frugiperda(J. E. Smith), invaded China in mid-December 2018; since then, it has become a great threat to Chinese agricultural production. Qinling Mountains–Huaihe River region(QM–HRR) is the transitional zone between northern and southern China, an important region for both corn and wheat production. Based on the actual occurrence of QM–HRR invaded by FAW in 2019, daily mean surface air temperature and nocturnal wind conditions at 925 hPa were examined, and migratory routes of FAW moths originated in QM–HRR were modeled by a forward-trajectory-analysis approach. The results indicated that migratory activities of FAW adults emerged in QM–HRR were initiated from late June. The moths from western QM–HRR, where has complex topographic terrain, mainly flied to Ningxia and Inner Mongolia before mid September. However, FAW moths from the eastern QM–HRR primarily engaged in high-altitude northward transport assisted by the prevailing southerly winds before mid August, and the North China Plain was identified as the main destination of FAW. Meanwhile, the migration trajectories of FAW moths had a possibility to reach the Northeast China Plain. From mid August, FAW moths in eastern QM–HRR largely migrated southward and returned to the Yangtze River Valley. This study provides detailed information on the occurrence and migration routes of FAW moths from QM–HRR and will be helpful for early warning and development of integrated pest management strategies for the control of this exotic insect pest.     

Spread of invasive migratory pest Spodoptera frugiperda and management practices throughout China

The fall armyworm(FAW), Spodoptera frugiperda(J. E. Smith)(Lepidoptera: Noctuidae), a notorious migratory pest native to tropical and subtropical America, invaded China in December 2018, then spread through 26 provinces(autonomous regions, municipalities) in 2019 and 27 in 2020, damaging 1.125 and 1.278 million hectares of crops, respectively. Maize was the most severely affected crop, although wheat and other plants were also ruined. Considering the biological characteristics, incidence regularity and migration patterns of the FAW populations, Chinese government implemented a regional control strategy and divided the areas infested with FAW into the annual breeding grounds in Southwest and South China, the transitional migration area in Jiangnan and Jianghuai and the key preventive area in the Huang-Huai-Hai region and North China. The National Agro-Tech Extension and Service Center constructed "the National Information Platform for the Prevention and Control of the Fall Armyworm" at the county level, which would entail people reporting and mapping the spread of fall armyworm. According to forecasting information, millions of extension workers and small-scale growers in entire country were rallied by local governments to fight the pest through comprehensive control tactics including chemical, physical, biological and ecological measures. Thanks to the joint prevention and control, the final loss of crops infested was controlled within 5% of the total in 2019 and 2020. This review also gives a discussion on existing problems and future management scenarios.     

Laboratory-based flight performance of the fall armyworm,Spodoptera frugiperda

In late 2018, the fall armyworm(FAW) Spodoptera frugiperda Smith(Lepidoptera: Noctuidae) made its arrival in China and its populations have since proliferated across most of eastern Asia. While S. frugiperda exhibits a considerable dispersal capacity and engages in long-distance migration, there's only scant information on the species' flight capability. Here, we empirically assessed S. frugiperda flight activity under varying climatic conditions using a flight mill. More specifically, under laboratory conditions, FAW exhibited superior flight performance at 20–25°C and 60–90% relative humidity(RH). When quantifying flight performance over five consecutive nights(i.e., 10 h/night), all flight parameters initially increased and then gradually dropped and FAW adults attained a total flight distance, duration and velocity equal to 63.73 km(48.42–94.12 km)(median, quartile range), 24.12 h(20.87–27.73 h) and 2.73 km h~(–1)(2.13–3.33 km h~(–1)), respectively. Our work constitutes a first comprehensive assessment of S. frugiperda flight performance and provides baseline information for future efforts to forecast spatiotemporal changes in its geographical distribution, movement patterns and invasion trajectories. Such can ultimately permit a timely and targeted deployment of area-wide pest management measures against this newly-invasive pest in China and across eastern Asia.     

Population occurrence of the fall armyworm,Spodoptera frugiperda(Lepidoptera: Noctuidae), in the winter season of China

Fall armyworm(FAW), Spodoptera frugiperda(J.E. Smith), has become an important pest in Chinese agricultural systems since its invasion on 11 December 2018. After its establishment, FAW months in the year-round breeding region have become the main source population migrating to other areas in China. Field investigations were conducted in tropical and subtropical regions to improve understanding of its year-round breeding area in China. The results showed that FAW larval density was significantly correlated with the seasonal temperature of the location surveyed. The FAW larvae maintained a high density in the tropical area and were frequently found in sites of the south subtropical region, but were absent from the north subtropical region and the northern part of the central subtropical region. These results indicated that FAW can reproduce annually in the tropical and south subtropical regions of China, including Hainan, Taiwan and the southern area of Fujian, Guangdong, Guangxi, Guizhou, and Yunnan. Hence, great effort should be made to monitor and control FAW in the year-round breeding region to suppress the population density of this area and to reduce migration of moths into northern parts of China. This study clarifies the occurrence area of the pest in winter in China and provides much valuable information for its population forecasting and management.     

Biology,invasion and management of the agricultural invader: Fall armyworm,Spodoptera frugiperda(Lepidoptera: Noctuidae)

The fall armyworm(FAW), Spodoptera frugiperda(J. E. Smith), is native to the Americas. It has rapidly invaded 47 African countries and 18 Asian countries since the first detection of invasion into Nigeria and Ghana in 2016. It is regarded as a super pest based on its host range(at least 353 host plants), its inherent ability to survive in a wide range of habitats, its strong migration ability, high fecundity, rapid development of resistance to insecticides/viruses and its gluttonous characteristics. The inherently superior biological characteristics of FAW contribute to its invasiveness. Integrated pest management(IPM) of FAW has relied on multiple applications of monitoring and scouting, agricultural control, chemical pesticides, viral insecticides, sex attractants, bio-control agents(parasitoids, predators and entomopathogens) and botanicals. Knowledge gaps remain to be filled to:(1) understand the invasive mechanisms of S. frugiperda;(2) understand how to prevent its further spread and(3) provide better management strategies. This review summarizes the biological characters of FAW, their association with its invasiveness and IPM strategies, which may provide further insights for future management.     

草地贪夜蛾在中国西北地区的迁飞路径

【背景】草地贪夜蛾（Spodoptera frugiperda）是联合国粮食及农业组织（FAO）全球预警的重大迁飞性农业害虫,2018年底入侵中国后对玉米等农作物生产构成了严重威胁。远距离迁飞习性是草地贪夜蛾区域性转移危害的生物学基础,其在东亚季风和印度季风流行的气候环境下,可形成纵贯中国东部地区和西部地区的迁飞路线。据连续两年全国普查,草地贪夜蛾沿西线迁飞路线逐代北侵的终点位于我国西北地区的宁夏全境和内蒙古阿拉善左旗。但目前,入侵中国西北地区的草地贪夜蛾的虫源地尚未明确,该害虫在整个中国西部地区的迁飞路径研究也鲜有报道。【目的】阐释驱动草地贪夜蛾迁入中国西北地区的关键气象动力因子,明确该地区首次入侵种群的虫源地所在,精准解析亚洲季风气候背景下草地贪夜蛾的迁飞路径,为建设监测预警体系和指导区域性治理提供依据。【方法】利用2019—2020年草地贪夜蛾在西北地区宁夏的成虫种群动态监测数据和气象资料,采用中尺度气象数值模拟、昆虫迁飞轨迹模型、地理信息系统等技术手段,分析草地贪夜蛾远距离迁飞的大气动力背景场,模拟计算草地贪夜蛾在中国西北地区连续1—3个夜晚的迁飞路径和回推轨迹落点。【结果】7—9月份出现的偏南夏季风是影响宁夏等西北地区草地贪夜蛾不断迁入的关键气象因子,其虫源主要来自于甘肃东南部、四川东部,其次为陕西西部,此外,重庆西南部、云南东北部和山西西部局部也可为西北地区提供一定的虫源。【结论】在偏南夏季风的主导作用下,草地贪夜蛾经由多个夜晚的连续迁飞可构成其在中国西部地区的主要迁飞路径。该路径源自缅甸,自南向北依次经由中国“云南-四川和重庆-陕西和甘肃-宁夏”,最北可达内蒙古。在7—9月份偏南风盛行期提前和风速偏强的气候条件下要加强西北部玉米种植区草地贪夜蛾的监测预警与防治工作。     

草地贪夜蛾对玉米的危害及防控对策

草地贪夜蛾Spodoptera frugiperda (J. E. Smith)是原产于美洲热带和亚热带地区的世界性农业害虫,具有迁徙能力强、繁衍速度快、危害程度重等特点。2019年1月,该虫第一次由东南亚侵入中国云南,在不到5个月的时间里扩散至江淮流域,并对我国主要经济作物—玉米造成了很大的威胁。文章综述了草地贪夜蛾的主要特征特点、对玉米的危害及主要的防控治理对策,为减轻草地贪夜蛾对我国玉米的危害提供依据。     

Ovipositional responses of Spodoptera frugiperda on host plants provide a basis for using Bt-transgenic maize as trap crop in China

Spodoptera frugiperda, the pest fall armyworm(FAW), is widespread in more than 100 countries. To date, planting insectresistant transgenic crops is one of the main control methods in its native countries. In this study we evaluated Bt-transgenic maize(Bt maize) and non-transgenic(conventional) maize and six other host plants in greenhouse pot experiments and field trials for oviposition preference by the Chinese populations of FAW. In laboratory trials, female moths preferred to oviposit on maize with no significant preference between conventional and Bt maize. However, after conventional and transgenic maize were exposed to FAW larvae and damaged, oviposition was significantly higher on transgenic maize than on the conventional one. Field trials showed that for plants less damaged at an early stage(seedling stage), oviposition of FAW on transgenic and conventional maize was significantly higher than that on wheat, sorghum, foxtail millet, peanut and soybean while showing no significant difference between transgenic or conventional maize. FAW adults mainly laid eggs on Bt maize, while the larval density and leaf damage rating or percentage of damaged plants were significantly lower than on conventional maize. Larval density and its damage on conventional maize were significantly higher than that on Bt maize and the other five hosts. Thus, maize is a highly preferred and suitable host for S. frugiperda feeding and ovipositing, and Bt maize can be used as trap crop to protect other crops.     

Flight activity promotes reproductive processes in the fall armyworm,Spodoptera frugiperda

The fall armyworm(FAW), Spodoptera frugiperda(Lepidoptera: Noctuidae), has invaded many countries in Africa and Asia since 2016, posing a major threat to world food security. Long-distance migration and strong reproductive ability form the biological basis of its rapid population expansion, but the relationship between the flight and reproduction of FAW has not been studied in depth. Here, an empirical assessment of this relationship in an invasive FAW population in China found that 1–3-day-old adults which had undergone 10-h tethered flights had a significantly shorter pre-oviposition period and greater oviposition synchronization, but did not show any differences in fecundity, oviposition period, mating percentage or other reproductive variables. Further studies on moths after 1.25–15-h tethered flights indicated that the reproductive process of adults could be fully triggered by flight activity longer than 2.5 h. Dissection of the reproductive organs also showed that tethered flight promoted ovarian and testicular development of FAW. These results show that appropriate moth flight activity significantly speeds up the reproductive process of FAW, which increases our knowledge on its migratory biology in relation to regional outbreaks.     

Case study on the first immigration of fall armyworm,Spodoptera frugiperda invading into China

The fall armyworm(FAW), Spodoptera frugiperda is one of the most damaging crop pests, and it has become major threat to the food security of many countries. In order to monitor possible invasion of this pest into China, a searchlight trap was established in March 2018 in western Yunnan Province, China, where it has served as the "first station" for many pests that have migrated from Myanmar to China. A number of suspected FAW moths were captured and identified by DNA sequencing. The results showed that the FAW moth was first captured on December 11 and formed its first immigration peak in mid-December 2018. DNA detection revealed that the early invading FAW population was the "corn-strain". The field survey indicated that the pest mainly colonized corn in Pu'er, Dehong and Baoshan areas. Migration trajectory simulation implied that the moths might have mainly come from the eastern area in the mid-latitude region of Myanmar(20–25°N, 94–100°E). This case study confirmed the first immigration of FAW into China, and will be helpful for guiding monitoring and management work to control this pest.     

Adult nutrition affects reproduction and flight performance of the invasive fall armyworm,Spodoptera frugiperda in China

Floral resources, such as carbohydrate-rich nectar or pollen, can bolster fitness and raise reproductive output of adult lepidopterans. Here, we used laboratory experiments to assess how those plant-derived foods impact adult fecundity, reproductive physiology and flight performance of an invasive strain of the fall armyworm, Spodoptera frugiperda(FAW; Lepidoptera: Noctuidae) in China. More specifically, supplementary feeding on bee pollen and honey enhanced FAW flight duration, testis size, ovarian development, longevity and adult fecundity. FAW adults attained the longest pre-oviposition(10.8 days) and oviposition period(6.8 days) and longevity(19.2 days) on 5% acacia honey. Upon access to 2.5% acacia honey and 2.5‰ pine pollen, S. frugiperda attained the highest mating rate(79.7%), fecundity(644.9 eggs/female) and egg hatching rate(82.3%). Feeding on honey further delayed decay of male testes, while ovarian development was enhanced when female moths were allowed access to 2.5% honey and 2.5‰ pine pollen. Upon feeding on 5% honey solution, S. frugiperda engaged in flight over the longest duration(9.5 h), distance(29.9 km) and speed(3.1 km h~(–1)). Honey had a comparatively greater effect on the above parameters than pollen. Our findings help decipher FAW invasion patterns and population dynamics, facilitate the development of nutritional attractants, and contribute to integrated pest management of this newly-invasive pest in eastern Asia.     

Population life tables for the invasive fall armyworm,Spodoptera frugiperda fed on major oil crops planted in China

The fall armyworm(FAW), Spodoptera frugiperda, is a newly invasive, widespread agricultural pest in China. Understanding the suitability of the main field crops in Chinese agricultural system as host for this polyphagous herbivore is especially important for making control strategy. Here, after FAWs were fed three important oil-bearing crops(oilseed rape, soybean and sunflower) planted in China and resultant population parameters were compared using the age-stage, two-sex life table method, survival of larvae on soybean was significantly lower than that on oilseed rape and sunflower. Developmental duration of larvae on soybean was also the longest(23.3 days). The highest pupation rate was recorded on sunflower. The highest pupal mass(0.19 g) was attained on oilseed rape, significantly higher than on the other host plants; the lowest mass was on soybean(0.15 g). On soybean, oilseed rape and sunflower, respectively, the average generation period was 42.21, 39.10 and 40.44 d; the intrinsic rate of increase(r) was 0.0844, 0.1041 and 0.1134; the finite rate of increase(λ) was 1.0881, 1.1098 and 1.1202. While the most suitable host plant overall was sunflower, S. frugiperda completed development and increased its population on all three host plants. Thus, soybean, oilseed rape and sunflower were all suitable for FAW, and population monitoring and management of FAW in these crops should be increased.     

草地贪夜蛾的综合防治

草地贪夜蛾Spodoptera frugiperda (J. E. Smith)属于入侵性农业害虫,其幼虫可以大量啃食玉米、高粱、水稻等多种农作物。草地贪夜蛾具有迁飞能力强、突发性、多产性、暴食性、杂食性、隐蔽性、钻蛀性、钻土性、夜出性、趋嫩性和独占性等特性,防治难度极大。对于入侵的暴发性害虫来说,化学防治依旧是目前能够在较短时间内控制其在我国大规模暴发的主要方法。虽然有很多昆虫都是草地贪夜蛾的天敌,但基本上没有足够用来防止作物受伤的措施。目前的防治措施主要包括物理防治、化学防治、生物防治,其中生物防治又可具体为植物源农药、性诱剂、生防菌、寄生蜂、寄生性天敌等。     

Fitness of fall armyworm,Spodoptera frugiperda to three solanaceous vegetables

The fall armyworm(FAW), Spodoptera frugiperda Smith(Lepidoptera: Noctuidae) is an important agricultural pest that invaded China in the middle of December 2018. As a polyphagous pest, FAW is identified as a serious threat to agricultural production and food security in China. Pepper(Capsicum annuum L.), tomato(Solanum lycopersicum Mill.) and eggplant(Solanum melongena L.) are three of dominant solanaceous vegetables of this country. To our knowledge, the effects of these plants on the performances of FAW have not been well studied. In this study we assessed the fitness of this pest to these three plants. Results showed that FAW can complete its life cycle when fed with tomato and pepper, but not on eggplant. The population parameters of FAW fed with maize(Zea mays L.) and the three solanaceous vegetables were compared using the age-stage, two-sex life table method. Developmental duration was significantly different in the larval stage, but not in the pupae stage. FAW fed with pepper had the longest pre-adult period(41.73 d) and the lightest pupal weight(0.1134 g); the survival rate was lower than FAW fed with tomato. Significant differences were observed in the mean fecundity of female, with the highest(943.95 eggs) laid by FAW fed with tomato. FAW had the shortest mean generation time(T), the highest intrinsic rate of increase(r) and finite rate of increase(λ) on maize, and the highest net reproductive rate(R_0) on tomato. Overall, FAW fitness on the three solanaceous vegetables was: tomatopeppereggplant. This study provides the foundation for further assessment of FAW risk to solanaceous vegetables and for establishing corresponding control strategies in China.     

西安地区夏玉米草地贪夜蛾发生及防控对策

草地贪夜蛾是一种跨境迁飞性重大害虫,具有寄主范围广、繁殖能力强、迁飞扩散快、为害程度重、防治难度大等特点。自2019年草地贪夜蛾入侵我国云南后,当年已扩散至包括陕西省在内的26个省份发生为害,发生面积5 013.36 hm,严重威胁粮食安全。2019年,西安各区县的玉米种植区草地贪夜蛾普遍发生,全市发生面积2 172.81 hm,占据全省的43.34%。文章对草地贪夜蛾在西安地区2019年的发生情况进行总结,旨在为今后草地贪夜蛾防控及可持续治理提供理论依据和技术支持。     

Larval diet affects development and reproduction of East Asian strain of the fall armyworm,Spodoptera frugiperda

In December 11, 2018, the fall armyworm(FAW), Spodoptera frugiperda invaded China and has since impacted local maize, sorghum and other crops. Here, we draw on laboratory experiments to show how different host crops(i.e., maize, sorghum, wheat and rice) and artificial diet affect larval growth and adult reproduction of one local FAW strain. Larval diet affected development duration, pupation rate, survival and emergence rate of pupae, and S. frugiperda adult fecundity. FAW attained the slowest larval development(19.4 days) on sorghum and the fastest(14.1 days) on artificial diet, with larvae attaining 99.6% survival on the latter food item. On rice, FAW larvae attained survival rate of 0.4% and were unable to pupate successfully. Pupation rate and pupal survival varied substantially between artificial diet and live plantlets at different phenological stages. Pupal weight was the highest(0.26 g) on artificial diet and the lowest(0.14 g) on sorghum, while FAW females reached the highest fecundity(699.7 eggs/female) on 2-leaf stage maize. Egg hatching rate equaled 93.6% on 4-or 5-leaf stage maize and 36.6% on artificial diet. FAW intrinsic rate of natural increase and the finite rate of increase varied between larval diets, reflecting how young maize leaves are the most suitable diet. Our findings can help to refine laboratory rearing protocols, devise population forecasting models or guide the deployment of ‘area-wide' integrated pest management(IPM) modules in FAW-invaded areas of China and other Asian countries.     

入侵重庆地区的草地贪夜蛾种群生物型鉴定

草地贪夜蛾是一种跨国迁飞性农业重大害虫,目前已在重庆的巫山、巫溪、云阳、北碚、潼南等地区出现并有迁飞的趋势.利用两个分子标记基因COI和Tpi对重庆地区采集的样本进行遗传特征比较,结果表明COI基因的亚型全为"水稻型",而Tpi基因的亚型既有"水稻型"又有"玉米型",说明入侵到重庆地区的草地贪夜蛾来源可能并不单一,暗示了重庆地区的草地贪夜蛾有可能存在分化为"水稻型"的风险.分析了入侵重庆地区的草地贪夜蛾相关的遗传特征,其结果不仅有助于认识草地贪夜蛾在中国大面积迁飞过程中发生的种群差异变化,也有利于农业部门根据重庆地区的地理环境属性,开展有针对性的监测防控工作.     

草地贪夜蛾入侵安徽

草地贪夜蛾Spodoptera frugiperda (Smith),是一种原产于美洲热带和亚热带的毁灭性农业害虫,2019年5月在安徽省黄山市黄山区首见该虫为害春玉米,对我省粮食生产造成威胁。本文就当前安徽省草地贪夜蛾发生情况、形态特征、危害特点、以及防治措施进行介绍。