Outbreak of an exotic flatid,Metcalfa pruinosa (Say) (Hemiptera: Flatidae), in the capital region of Korea

The citrus flatid planthopper, Metcalfa pruinosa (Say, 1830) (Hemiptera: Flatidae), has a native distribution in eastern North America, It has recently invaded Italy in 1979 and has since spread to other European countries. In 2009, Metcalfa pruinosa was discovered in Seoul and the Gyeonggi Province, Republic of Korea. This is the first record in the eastern part of Palaearctic. One year after its discovery, in July 2010, we found significant populations and serious damage on many deciduous forest trees, ornamental trees, and agricultural crops in central regions of the Korean Peninsula. In this paper, we report the status of the outbreak and discuss the biology, morphological characters, distribution, host plants, and the importance of M. pruinosa as a potential insect pest in the Korean Peninsula.     

Biological and molecular evidence for the transmission of aster yellows phytoplasma to French marigold (Tagetes patula) by the flatid planthopper Metcalfa pruinosa

Aster yellows (AY) phytoplasmas (Candidatus Phytoplasma asteris) are associated with a number of plant diseases throughout the world. Several insect vectors are responsible for spreading AY diseases resulting in wide distribution and low host specificity. Because the role of sucking insects as vectors of phytoplasmas is widely documented, and the citrus flatid planthopper Metcalfa pruinosa is a phloem feeder, it has been incriminated as a possible vector of phytoplasmas. However, its ability to transfer phytoplasma has not been confirmed. The present work shows that M. pruinosa (Hemiptera: Flatidae), a polyphagous planthopper, is able to vector Ca. P. asteris to French marigold (Tagetes patula). Transmission experiments were conducted in 2017 and 2018 in central Hungary by two approaches: (a) AY-infected M. pruinosa were collected from an area with severe incidence of the disease on T. patula and caged on test plants for an inoculation-access period of 2 weeks, and (b) presumably phytoplasma-free insects were collected from apparently healthy grapevines (Vitis vinifera L.) and fed on AY-infected T. patula plants for 2 weeks prior to being caged on test plants. AY disease symptoms developed on 4 out of 10 and 10 out of 15 test plants, respectively. All phytoplasma-positive marigold and M. pruinosa samples showed identical RFLP patterns and shared 100% 16S rDNA sequence identity with each other and with the aster yellows phytoplasma strain AJ33 (GenBank accession number MK992774). These results indicated that the phytoplasma belonged to the phytoplasma subgroup 16SrI-B Ca. P. asteris. Therefore, the work presented here provides experimental evidence that M. pruinosa is a vector of a 16SrI-B subgroup phytoplasma to T. patula.     

Plant penetration activities by the flatid planthopper Metcalfa pruinosa (Hemiptera: Fulgoroidea): an electrical penetration graph‐histology analysis

The citrus flatid planthopper, Metcalfa pruinosa Say (Hemiptera: Flatidae), is a very polyphagous native insect in North America and currently a serious pest in Europe and South Korea. To understand the feeding behaviour of M. pruinosa, stylet penetration behaviour of M. pruinosa was investigated with an electrical penetration graph (EPG) system. This study reports seven EPG waveforms related to M. pruinosa feeding behaviour: np (no stylet penetration), Mp1 (initiation of stylet penetration), Mp2 (stylet movement and salivation), Mp4 (phloem feeding), Mp4‐H (honeydew excretion), Mp5 (xylem feeding) and Mp6 (unknown). To determine respective feeding behaviour related to the Mp4 and Mp5 waveforms, stylets were cut with a laser beam, and the location of the stylet tip within plant tissue was examined. We found plant sap was exuded from the severed stylets only when the Mp4 waveform was observed, suggesting phloem sap ingestion. The stylet tip was located in the xylem region, indicating xylem‐feeding activity, when the Mp5 waveform was observed. The analysis of 24 different EPG parameters suggests that M. pruinosa stylets reached the vascular bundle of a plant within ca. 5 min and spend ca. 70% of the time feeding on xylem and phloem feeding. This is the first study that reports seven distinctive EPG waveforms with respect to the feeding behaviour of M. pruinosa which could help determine host specificity and host plant susceptibility.     

Predicting potential current distribution of Lycorma delicatula (Hemiptera: Fulgoridae) using MaxEnt model in South Korea

Lycorma delicatula (Hemiptera: Fulgoridae) is an invasive insect in Korea which causes plant damages by sucking and sooty molds. Lycorma delicatula was first detected in South Korea in 2004, where its introduction and spreading possibly were affected by human activity-related factors. Here, we used MaxEnt to describe current distribution of L. delicatula in Korea and tried to find out the impact of human influences on distribution. We used 143 sites of occurrence data, 19 bioclimatic variables, duration of temperature below −11 °C, average daily minimum temperature in January, cumulative thermal unit variable, the distribution of grape orchard variable and human footprint to create models. These models were estimated by two sets of 24 candidates with feature combinations and regularization multipliers. In addition, these two sets were created as models with and without footprint to assess human influence on distribution. Model selection for optimal model was performed by selecting a model with a lowest sum of each rank in small sample-size corrected Akaike’s information criterion and difference between training and test AUC. Model of LQ10 parameter combinations was selected as optimal models for both model sets. Consequently, both of distribution maps from these models showed similar patterns of presence probability for L. delicatula. Both models expected that low altitude regions were relatively more suitable for L. delicatula than mountain areas in Korea. Footprint might be limited for the distribution and L. delicatula might already occupy most of available habitats. Human-related factors might contribute to spread of L. delicatula to uninfected areas.     

Multigene phylogeny and haplotype analysis reveals predominance of oomycetous fungus,Phytophthora meadii (McRae) associated with fruit rot disease of arecanut in India

An oomycetous fungus Phytophthora causing fruit rot is the most devastating disease of arecanut in different agro-climatic zones of Karnataka with varied climatic profiles. The main aim of this investigation was to characterize the geo-distant Phytophthora populations infecting arecanut using robust morphological, multi-gene phylogeny and haplotype analysis. A total of 48 geo-distant fruit rot infected samples were collected during the South-West monsoon of 2017–19. Pure culture of the suspected pathogen was isolated from the infected nuts and pathogenic ability was confirmed and characterized. Colony morphology revealed typical whitish mycelium with stellate or petalloid pattern and appearance with torulose hyphae. Sporangia were caducous, semipapillate or papillate, globose, ellipsoid or ovoid-obpyriform in shape and sporangiophores were irregularly branched or simple sympodial in nature. Subsequent multi-gene phylogeny (ITS, β-tub, TEF-1α and Cox-II) and sequence analysis confirmed the identity of oomycete as Phytophthora meadii which is predominant across the regions studied. We identified 49 haplotypes representing the higher haplotype diversity with varying relative haplotype frequency. Comprehensive study confirmed the existence of substantial variability among geo-distant populations (n = 48) of P. meadii. The knowledge on population dynamics of the pathogen causing fruit rot of arecanut generated from this investigation would aid in developing appropriate disease management strategies to curtail its further occurrence and spread in arecanut ecosystem.     

世界毁灭性检疫害虫番茄潜叶蛾的生物生态学及危害与控制

番茄潜叶蛾原产于南美洲的秘鲁,严重危害多种茄科作物,是最具毁灭性的世界检疫性害虫,严重发生地块番茄减产80%~100%。该虫主要借助农产品的贸易活动进行远距离传播扩散。截至2017年,番茄潜叶蛾已在全世界的85个国家和地区发生(以及在22个国家和地区疑似发生)。目前,我国尚没有该虫分布,然而其一旦入侵,将对我国的番茄和马铃薯产业构成巨大威胁。从番茄潜叶蛾寄主范围、危害特点及造成的经济损失、生物生态学习性、地理分布和传播扩散途径、防控措施等方面进行概述,以期为该虫的有效防范提供参考。     

Spatio-temporal distribution of Ricania shantungensis (Hemiptera: Ricaniidae) in chestnut fields: Implications for site-specific management

Chestnut is one of most important forest products in Korea. However, Ricania shantungensis, an invasive species, has been increasingly causing economic damage to chestnut. To increase management efficiency of R. shantungensis in chestnut fields, its spatio-temporal distributions were analyzed using spatial analysis by distance indices (SADIE) and semivariogram. Experiments were conducted by observing selected, marked, and geo-referenced trees in two commercial chestnut fields for each developmental stage (i.e., spring egg, nymph, adult, and fall egg) of R. shantungensis from 2017 to 2019. Spatial distributions of R. shantungensis were statistically (P < 0.05) aggregated except for its nymphal stage. Spatial associations of its distributions also showed the statistically (P < 0.05) positive associations regardless of years or developmental stages except for the spatial relationship between egg and nymphal stages. These results indicated that site-specific management for R. shantungensis would be applicable. In application of site-specific management for R. shantungensis, the distance of spatial dependence for nymphs, 60 m, should be considered to minimize reoccurrence possibility and additional samplings. Incorporating site-specific management into pest control program of R. shantungensis could increase its control efficiency in chestnut fields.     

Review of the Genus Myrmica in Korea (Hymenptera: Formicidae)

The paper contains a taxonomie review of 12 species of the genus Myrmica occurring in Korea. The following species are under discussion: M. angulinodis Ruzsky, M. carinata Kupyanskaya, M. hyungokae Elmes, M. jessensis Forel, M. kasczenkoi Ruzsky, M. kotokui Forel, M. koreana Elmes, M kurokii Forel, M. lobicornis Nylander, M. ruginodis Nylander, M silvestrii Wheeler, M. sulcinodis Nylander. Twelve species of the genus Myrmica are recognized from Korea, these are revised and a key to their identification is provided. Five species, M. cadusa, M. incurvata, M. saphoshikovi, M. scabrinodis, and M. yoshiokai are questionable to be distributed in Korea, as I have not found any specimens preserved in Korea. The SEM photo and illustrations of the worker of each species are provided.     

CLIMEX simulated predictions of the potential distribution of Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) considering the northern boundary: With special emphasis on Jeju,Korea

The global climate change has led to the distribution of Bactrocera dorsalis from its origin in Southeast Asia to mainland China, which is situated at latitudes similar to Jeju Island, South Korea. In order to evaluate the establishment of B. dorsalis in Korea, it is necessary to determine the climate suitability. When the parameters of CLIMEX model were adjusted to include Wuhan region in China as the northern limit, the model outputs explained well the distribution and occurrence characteristics (namely the annual number of generations) of B. dorsalis in major distribution regions in the world. In IPCC/RCP 8.5 scenario, the establishment of B. dorsalis was suitable to a small part of Jeju Island in the 2010s. The range might gradually spread until the 2090s, with many parts of the lowlands of Jeju Island being marginal to optimal in suitability. The present CLIMEX model with parameters that adjusted to known northern limit will be useful for annual risk mapping to facilitate a monitoring system for early detection of a possible founder population of B. dorsalis in Jeju, South Korea.     

Competitive habitat utilization in the damselfly,Mnais nawai (Zygoptera: Calopterygidae) coexisting with a related species,Mnais pruinosa

Reproductive behaviors related to habitat utilization were studied in males of the damsefly,Mnais nawai, which has two male forms, territorial orange-winged males (nawai) and non-territorial pale-orange-winged males (sahoi), at the upper part of a mountain stream where they partiallycoexist with a related species,Mnais pruinosa, which also has two male forms, territorial orange-winged males (esakii) and non-territorial hyaline-winged males (strigata). These two species showed parapatric distribution; the lower part of the stream was occupied byM. nawai, and the upper part byM. pruinosa. In the present study, cross-matings occurred between bothMnais species, although normal intraspecific matings occurred more frequently than cross-matings. Territorial males of both species copulated with conspecific females that entered their territory and guarded the ovipositing females, probably to avoid sperm displacement resulting from subsequent copulations. Severe competition for oviposition sites by territorial males even occurred between the two species. On the other hand, non-territorial males of both species have alternative mating strategies (including several tactics such as sneaking, takeover and interception). The possible benefits from conflict among territorial males of both species is discussed.     

Development of Sex Pheromone Trap for Monitoring Matsumuraeses falcana (Walshingham) (Lepidoptera: Tortricidae)

This study was carried out to find out an effective formulation for monitoring the soybean podborer, Matsumuraeses falcana (Walshingham), using its sex pheromone in Korea. A 1 : 1 : 1 mixture of (E)-8-dodecenyl acetate (E 8–12: Ac), (E,E)-8,10-dodecadienyl acetate (E 8 E 10–12: Ac) and (E,Z)-7,9-dodecadienyl acetate (E 7 Z 9–12: Ac) was an effective composition in monitoring M. falcana. A binary mixture of E 8–12: Ac and E 8 E 10–12: Ac was essential to attract M. falcana male moths. E 7 Z 9–12: Ac showed a synergistic effect on the attractiveness of 1 : 1 mixture of E 8–12: Ac and E 8 E 10–12: Ac. These chemicals alone showed little or no attractiveness to M. falcana. The sticky-typed wing trap baited with sex pheromone captured significantly more males than the water-pan trap. Trap catches within a soybean field were much more than those in perimeter trap. The trap-baited with sex pheromone gave a distinct fluctuation pattern of male catches throughout the flight seasons. Therefore, it can be expected that pheromone-baited trap provides a simple tool for monitoring the soybean podborer, M. falcana in Korea.     

Anthropogenic and Ecological Drivers of Amphibian Disease (Ranavirosis)

Ranaviruses are causing mass amphibian die-offs in North America, Europe and Asia, and have been implicated in the decline of common frog (Rana temporaria) populations in the UK. Despite this, we have very little understanding of the environmental drivers of disease occurrence and prevalence. Using a long term (1992-2000) dataset of public reports of amphibian mortalities, we assess a set of potential predictors of the occurrence and prevalence of Ranavirus-consistent common frog mortality events in Britain. We reveal the influence of biotic and abiotic drivers of this disease, with many of these abiotic characteristics being anthropogenic. Whilst controlling for the geographic distribution of mortality events, disease prevalence increases with increasing frog population density, presence of fish and wild newts, increasing pond depth and the use of garden chemicals. The presence of an alternative host reduces prevalence, potentially indicating a dilution effect. Ranavirosis occurrence is associated with the presence of toads, an urban setting and the use of fish care products, providing insight into the causes of emergence of disease. Links between occurrence, prevalence, pond characteristics and garden management practices provides useful management implications for reducing the impacts of Ranavirus in the wild.     

Occurrence model of first instars of Ricania shantungensis (Hemiptera: Ricaniidae)

An invasive planthopper, Ricania shantungensis, is an important pest in agriculture and forestry in Korea. Best target stage for insecticide application is known to be newly hatched first instar. Thus, the objective of the present study was to predict the occurrence of first instars of R. shantungensis. Effects of temperature on development and survival of R. shantungensis eggs were examined at seven constant temperatures (12.4, 16.4, 20.4, 24.8, 28.3, 32.4, and 36.9 °C). Development and survival of R. shantungensis eggs were quantitatively described by applying empirical models as a function of temperature over a wide thermal range. Lower developmental threshold, thermal constant, optimal developmental temperature, and upper developmental threshold were estimated to be 12.1 °C, 202 DD, 31 °C, and 36.9 °C, respectively. Survivorship was the highest at 23.3 °C. The models well predicted timing of field occurrences at three sites (Buyeo, Gwangyang, and Habcheon) in Korea. Therefore, results of this study would increase the prediction accuracy of R. shantungensis occurrence and management efficiency of R. shantungensis.     

Predicting Microstegium vimineum invasion in natural plant communities of the southern Blue Ridge Mountains, USA

Shade-tolerant non-native invasive plant species may make deep incursions into natural plant communities, but detecting such species is challenging because occurrences are often sparse. We developed Bayesian models of the distribution of Microstegium vimineum in natural plant communities of the southern Blue Ridge Mountains, USA to address three objectives: (1) to assess local and landscape factors that influence the probability of presence of M. vimineum; (2) to quantify the spatial covariance error structure in occurrence that was not accounted for by the environmental variables; and (3) to synthesize our results with previous findings to make inference on the spatial attributes of the invasion process. Natural plant communities surrounded by areas with high human activity and low forest cover were at highest risk of M. vimineum invasion. The probability of M. vimineum presence also increased with increasing native species richness and soil pH, and decreasing basal area of ericaceous shrubs. After accounting for environmental covariates, evaluation of the spatial covariance error structure revealed that M. vimineum is invading the landscape by a hierarchical process. Infrequent long-distance dispersal events result in new nascent sub-populations that then spread via intermediate- and short-distance dispersal, resulting in 3-km spatial aggregation pattern of sub-populations. Containment or minimisation of its impact on native plant communities will be contingent on understanding how M. vimineum can be prevented from colonizing new suitable habitats. The hierarchical invasion process proposed here provides a framework to organise and focus research and management efforts.     

Host range testing of the nearctic beneficial parasitoid <Emphasis Type="Italic">Neodryinus typhlocybae</Emphasis>

Neodryinus typhlocybae (Ashmead) (Hymenoptera: Dryinidae) is a natural enemy of the planthopper Metcalfa pruinosa (Say) (Hemiptera: Flatidae), introduced from North America into Europe and regionally established as a pest species. Prior to possible utilization of the parasitoid as a biocontrol agent in Austria, its potential negative impacts on eight native plant- and leaf-hopper species were examined in the laboratory. Non-target species were selected according to the following criteria (a) occurrence in Austria, (b) close phylogenetic relationship with M. pruinosa, (c) larvae free-living and surface-dwelling, (d) phenology, (e) larval size, (f) ecological similarity with M. pruinosa and (g) availability of sufficient numbers of individuals. The Auchenorrhyncha species Issus coleoptratus (Fabricius), Chloriona smaragdula (Stål), Conomelus anceps (Germar), Alebra wahlbergi (Boheman), Empoasca sp., Idiocerus stigmaticalis (Lewis), Macrosteles septemnotatus (Fallén) and Japananus hyalinus (Osborn) were chosen for testing. Larvae from both the target and the non-target species were offered separately to N. typhlocybae females in no-choice laboratory tests and all attacks, instances of host feeding and parasitizations were recorded. No non-target species was attacked, fed upon or parasitized by N. typhlocybae, whereas M. pruinosa was attacked frequently. This study supports the assumption that the host range of N. typhlocybae is restricted to Flatidae, of which only the introduced species occurs in Austria. Direct negative effects on other Auchenorryncha species in Austria are therefore unlikely to occur.     

The spread of non-native <Emphasis Type="Italic">Cepaea nemoralis</Emphasis> and <Emphasis Type="Italic">Monacha cartusiana</Emphasis> (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants

The aim of our study is to describe and visualise the spread of two non-indigenous land snail species Cepaea nemoralis and Monacha cartusiana in the Czech Republic during more than 100 years period. Several factors play an important role in changes of the distribution of these species: ecological (climate change), ethological (passive dispersal potencial) and economic (increasing traffic as a vector of spreading). The spreading of M. cartusiana has a rapidly increasing trend. More than half sites in the Czech Republic were colonised by this species in 2000–2010. While the spread of C. nemoralis has been continuous during the last century, the rapid range extension was recorded in the last two decades.     

Changes in spatial and temporal distributions of Monochamus beetles along the fire severity in burned Pinus densiflora forests

Wildfire is an important disturbance factor in forest ecosystem and could affect the distribution of insects. Because Monochamus beetles are known vector insects of pine wood nematodes in Korea and burnt pine trees can be used as oviposition habitats for Monochamus beetles, we investigated changes in the spatial (among fire severity classes) and temporal (between years) distributions of M. alternatus and M. saltuarius. From late May 2017 to early October 2018, four multi-funnel traps with pheromone lure were installed in each fire severity class (unburned, light, light-moderate edge, moderate-severe edge, and severe). All traps were replaced fortnightly from May to October for two years. The spatial distribution of the two Monochamus beetles was significantly affected by fire severity. In particular, M. alternatus showed higher catches in the severely burned sites and responded positively to the fire. A similar number of M. saltuarius individuals were observed between 2017 and 2018, while more M. alternatus were caught in 2018 compared to 2017. Consequently, it was confirmed that the dead wood of P. densiflora caused by the wildfire had a great influence on the density of insects, especially M. alternatus.     

Discovery of an egg parasitoid of Lycorma delicatula (Hemiptera: Fulgoridae) an invasive species in South Korea

Anastatus sp. (Hymenoptera: Eupelmidae) is reported as the first chalcidoid wasp which parasitizes eggs of Lycorma delicatula, an invasive species spreading rapidly in South Korea. The wasp appears to be a solitary endoparasitoid. The present paper also describes its oviposition and host feeding behaviour.     

Phylogeny and divergence time estimation of Coreoleuciscus splendidus populations (Teleostei: Cypriniformes) endemic to Korea based on complete mitogenome sequences

We determined the complete mitogenome sequences for four Coreoleuciscus splendidus strains (Teleostei: Cypriniformes) inhabiting the Han and Geum river drainages in the West Korea Subdistrict, and the Seomjin and Nakdong river drainages in the South Korea Subdistrict in the Korean peninsula. The gene contents and arrangement of C. splendidus were homogeneous to those of typical vertebrates. Phylogenetic analysis with the partitioned nucleotide matrix of concatenated mitochondrial genes revealed that C. splendidus formed a monophyletic group with gobionine species in the cyprinid lineage. The four river strains separated into two phylogenetically distinct groups, which were in accordance with the biogeographical distribution pattern in Korea according to subdistrict. The estimated divergence time among western and southern subdistrict populations was 37.9 ± 6.5 million years ago (Mya), whereas those between the two river strains in each subdistrict were estimated to be 3.2?C3.4 Mya. The presence of such distinct historical lineages has great implications for biogeography of ichthyofauna in Korea and future management and conservation plans of C. splendidus populations.