Current and future distribution of Metcalfa pruinosa (Say) (Hemiptera: Flatidae) in Korea: Reasoning of fast spreading

Metcalfa pruinosa (Say) has rapidly established and spread through Korea since its first reported occurrence in 2009, causing serious economic damage to various agricultural crops. Several statistical modeling methods and MaxEnt software were used with spatio-temporal occurrence information in Korea, in order to verify fast expansion triggered by human-mediated factor, and predict potential distribution and its determinants of M. pruinosa for efficient management. Results showed that the estimated spread rate and pattern of M. pruinosa in Korea were significantly different from a continuous spreading pattern. In early spread phase in 2013, traffic volume, one of human mediated factors, was the most important environmental factor. Traffic volume of the relative pre-occupied habitat in 2013 was also significantly (P < .05) higher than the one lately occupied in 2017. However, the mean temperature in the warmest quarter became the most important environmental factor in current invasion phase of 2017. According to these results, traffic volume was strongly proposed as a main reason of rapid spreading of M. pruinosa in Korea, causing long-distance dispersal in stratified spread mechanism. The predicted current and future distribution of M. pruinosa can be used to prepare management plans and increase management efficiency for M. pruinosa in Korea.     

Prediction of global geographic distribution of Metcalfa pruinosa using CLIMEX

Globalization has changed the habitats of various species, resulting in harmful pest invasion. Among these pests, Metcalfa pruinosa has caused worldwide economic and hygienic damage in both urban and agricultural/forested areas. It has been reported that prediction of pest distribution is key to the management of pest prevention. Hence, this study aimed to predict the potential geographic distribution of M. pruinosa under the current climate and under a climate change scenario. CLIMEX, modeling software that analyzes the habitat suitability of a target species based on comprehensive climatic and physiological data, was used mainly to establish a map of predictive distribution of M. pruinosa at present and in the future. Based on our simulations, we predict that M. pruinosa will tend to extend its distribution northward in North America and Europe. We conclude that climate change could result in M. pruinosa invasion in a northward direction, suggesting the need for a thorough system of control and prevention.     

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.     

Phylogeography of Fischer’s blue,Tongeia fischeri,in Japan: Evidence for introgressive hybridization

The widespread lycaenid butterfly Tongeia fischeri is distributed from eastern Europe to northeastern Asia and represented by three geographically isolated populations in Japan. In order to clarify the phylogeographic history of the species, we used sequences of three mitochondrial (COI, Cyt b and ND5) and two nuclear (Rpl5 and Ldh) genes of 207 individuals collected from 55 sites throughout Japan and five sites on the Asian continent. Phylogenetic trees and the median-joining network revealed six evolutionary mitochondrial haplotype clades, which corresponded to the geographic distribution of the species. Common ancestors of Japanese T. fischeri might have come to Japan during the mid-Pleistocene by multiple dispersals of continental populations, probably via a land bridge or narrow channel between western Japan and the Korean Peninsula. The geographical patterns of variation of mitochondrial and nuclear markers are discordant in northeastern Kyushu, possibly as a result of introgressive hybridization during the ancient contact between the Kyushu and Shikoku populations in the last glacial maximum. The phylogeographic pattern of T. fischeri in Japan are probably related to the geological history, Pleistocene climatic oscillations and distribution of the host plant.     

Phylogeography and genetic diversity of East Asian Neolitsea sericea (Lauraceae) based on variations in chloroplast DNA sequences

Neolitsea sericea is an evergreen broad leaved tree in the warm-temperate regions of East Asia. This area is a hotspot for plant species richness and endemism caused by dynamic changes in land configuration during the Quaternary. However, the historical migration of such evergreen tree species is still poorly understood. In an attempt to reconstruct the phylogeographic history of N. sericea during the Quaternary, we identified the chloroplast DNA haplotypes of 287 individuals from 33 populations covering almost all of its geographic range. Analyses were based on sequence data from the trnL–F, psbC–trnS, and rps16 regions. Nine haplotypes were identified. The majority included ancestral types in the southwestern part of the main islands of Japan, with other region-specific haplotypes being found in populations on the Korean Peninsula, Taiwan (Isl. Lanyu), and elsewhere in Japan. A statistical parsimony network revealed two lineages derived from Japanese main islands. One was represented on the Korean Peninsula, the other on Isl. Lanyu. The current distribution of N. sericea has been shaped by colonization via land bridges. During the glacial periods, two primary, but separate migration routes were followed—from the southwestern part of the Japanese main islands to either the Korean Peninsula or Taiwan. In addition, we believe the Zhoushan populations were shaped by post-glacial processes through an ECS land bridge (East China Sea basin) from northern refugia that existed during the late Pleistocene.     

Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae): Genomic Comparison and Phylogenetic Inference in Fulgoroidea

The complete mitochondrial genomes (mitogenomes) of two DNA barcode-defined haplotypes of Metcalfa pruinosa and one of Salurnis marginella (Hemiptera: Flatidae) were sequenced and compared to those of other Fulgoroidea species. Furthermore, the mitogenome sequences were used to reconstruct phylogenetic relationships among fulgoroid families. The three mitogenomes, including that of the available species of Flatidae, commonly possessed distinctive structures in the 1702–1836 bp A+T-rich region, such as two repeat regions at each end and a large centered nonrepeat region. All members of the superfamily Fulgoroidea, including the Flatidae, consistently possessed a motiflike sequence (TAGTA) at the ND1 and trnS₂ junction. The phylogenetic analyses consistently recovered the familial relationships of (((((Ricaniidae + Issidae) + Flatidae) + Fulgoridae) + Achilidae) + Derbidae) in the amino acid-based analysis, with the placement of Cixiidae and Delphacidae as the earliest-derived lineages of fulgoroid families, whereas the monophyly of Delphacidae was not congruent between tree-constructing algorithms.     

Predicting potential distribution of chestnut phylloxerid (Hemiptera: Phylloxeridae) based on GARP and Maxent ecological niche models

The chestnut phylloxerid, Moritziella castaneivora, has been recently recorded as a forest pest in China. It heavily damaged chestnut trees and has caused serious economic losses in some main chestnut production areas. In order to effectively monitor and manage this pest, it is necessary to investigate its potential geographical distribution worldwide. In this study, we used two ecological niche models, Genetic Algorithm for Rule‐set Production (GARP) and Maximum Entropy (Maxent), along with the geographical distribution of the host plants, Japanese chestnut (Castanea crenata) and Chinese chestnut (Castanea mollissima), to predict the potential geographical distribution of M. castaneivora. The results suggested that the suitable distribution areas based on GARP were general consistent with those based on Maxent, but GARP predicted distribution areas that extended more in size than did Maxent. The results also indicated that the suitable areas for chestnut phylloxerid infestations were mainly restricted to Northeast China (northern Liaoning), East China (southern Shandong, northern Jiangsu and western Anhui), North China (southern Hebei, Beijing and Tianjin), Central China (eastern Hubei and southern Henan), Japan (Kinki, Shikoku and Tohoku) and most parts of the Korean Peninsula. In addition, some provinces of central and western China were predicted to have low suitability or unsuitable areas (e.g. Xinjiang, Qinghai and Tibet). A jackknife test in Maxent showed that the average precipitation in July was the most important environmental variable affecting the distribution of this pest species. Consequently, the study suggests several reasonable regulations and management strategies for avoiding the introduction or invasion of this high‐risk chestnut pest to these potentially suitable areas.     

Population genetic structure of endangered Mongolian racerunner (Eremias argus) from the Korean Peninsula

The Mongolian racerunner (Eremias argus) is a small lacertid lizard species, and its distribution range encompasses the Korean Peninsula, Mongolia, China and Russia. Eremias argus is widespread, but populations on the Korean Peninsula are small and declining, provoking concerns that genetic diversity is being lost. This species is currently listed under the Protection of Wild Fauna and Flora Act in South Korea. In this study, nine novel microsatellites for E. argus were developed with a biotin-enrichment method and used to understand its population genetic structure and delineate conservation units on the Korean Peninsula. Overall, low intrapopulation genetic diversity was observed (mean number of alleles per locus = 2.463; mean H _E = 0.398) from 10 populations investigated (n = 110). Two populations (among five with n≥ 10) showed an excess of heterozygosity expected under HWE relative to that expected at mutation-drift equilibrium, indicating severe reduction in population sizes. With only a few exceptions, the overall genetic differentiation among populations was substantial with the high levels of pairwise-F _ST (0.006–0.746) and -R _ST (0.034–0.940) values. The results of Bayesian STRUCTURE analysis showed that E. argus populations on the Korean Peninsula were most likely partitioned into three genetic clusters. Taken all together, such low levels of gene flow and strong genetic structuring have critical implications for the conservation of this endangered species and its management.     

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.     

The Johnston's organ of three homopteran species: A comparative ultrastructural study

A transmission electron-microscopy study has been carried out on the pedicel of three homopteran species, with particular focus on the leafhopper Scaphoideus titanus Ball. The two other species, the planthoppers Hyalesthes obsoletus Signoret and Metcalfa pruinosa Say, were investigated in order to compare the ultrastructure of the Johnston's organ (JO) among representatives of the Auchenorrhyncha group. The results showed the presence of a well developed JO located within the pedicel. Depending on the species the JO is made of 25 up to 72 scolopidia arranged in a coronal array. Each scolopidium is connective, heterodynal, amphinematic and hosts three structurally dissimilar sensory neurons. Two of them have a type 1 ciliary segment while the third bears a type 2 cilium. The type 2 dendrite tip is associated with a tubular cap and is longer than the others, ending into the cuticle at the base of the flagellum. Other scolopidia with one or two neurons were found in S. titanus, forming an accessory organ. The presence of such a well developed mechanosensory apparatus is discussed in relation with the lifestyle of the three species.     

New Record of the Family Lasiochilidae Carayon, 1972 (Hemiptera: Cimicoidea) from the Korean Peninsula,with Re-Description of Lasiochilus (Dilasia) japonicusfrom Jeju Island

Herein, Lasiochilus (Dilasia) japonicus Hiura 1967 is recognized for the first time on the Korean Peninsula. This insect is a member of the Family Lasiochilidae, which is new to the Korean Peninsula. Both female and male adults are re-described on the basis of specimens obtained from Jeju Island, Korea.     

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.     

Comparative genomics and selection analysis of Yeonsan Ogye black chicken with whole-genome sequencing

Yeonsan Ogye (OGYE; Gallus gallus domesticus) is a rare indigenous chicken breed that inhabits the Korean Peninsula. This breed has completely black coloring, including plumage, skin, eyes, beak, and internal organs. Despite these unique morphological characteristics, the population of OGYE has declined without in-depth research into their genome research. Therefore, this study aimed to compare the whole genome of OGYE to 12 other chicken populations, including ancestral breed, commercial breeds, Chinese indigenous breeds, and Korean native chickens. We focused on revealing the selection signature of OGYE, which has occurred through environmental pressures in the Korean Peninsula. Genome-wide selection analysis has identified local adaptation traits, such as egg development, that contribute to fetal viability and innate immune response to prevent viral and microbes infection in OGYE. In particular, SPP1 (Secreted Phosphoprotein 1), HSP90AA1 (Heat Shock Protein 90 Alpha Family Class A Member 1), and P2RX4 (Purinergic Receptor P2X 4) could have considerable involvement in egg development and RNASEL (Ribonuclease L), BRIP1 (BRCA1 Interacting Protein C-terminal Helicase 1), and TLR4 (Toll-Like Receptor 4) are crucial for the determination of the innate immune response. This study revealed the unique genetic diversity of OGYE at the genome-wide level. Furthermore, we emphasized the sustainable management of genetic resources and formulated breeding strategies for livestock on the Korean Peninsula.     

CYTOLOGICAL PROOF OF APOMIXIS IN CRATAEGUS (ROSACEAE)

An abundance of circumstantial evidence has led earlier workers to believe that apomixis was a common phenomenon in North American Crataegus. However, direct proof was lacking. In this paper, working with accessions of Crataegus pruinosa (Wendl.) K. Koch, sens. lat., a common and rather variable species of eastern North America, we present cytological proof of apomixis. This paper demonstrates the existence of apospory and documents the development of the aposporous embryo sac. Pseudogamy is inferred.     

Two species of the subfamily Olethreutinae (Lepidoptera: Tortricidae) new to the Korean Peninsula

Two species of the subfamily Olethreutinae, Epinotia maculana Fabricius and Zeiraphera lariciana Kawabe, are reported as new to the Korean Peninsula. Photographs of adults and genitalia are provided, with brief comments on the distribution and biology.     

Speciation processes within the Korean Peninsula based on the mites from the family Zerconidae (Acari: Mesostigmata)

The mesostigmatid mites speciation of the family Zerconidae in the Korean Peninsula was analysed in relation to the range of this group in the world. The Korean Peninsula is located on the edge of the Holarctic and is characterised by a uniquely diversified climate shaping the longitudinally changing flora. At present, there are altogether 40 genera included in the family Zerconidae. So far, in the Korean Peninsula there have been recognised 18 species of the Zerconidae belonging to 11 genera. The endemic genera within the area include: Metazercon, Eurozer-con, Xenozercon, Koreozercon, Kaikiozercon, Aquilo-nozercon, and cf. Mesozercon gen. nov. A very high level of endemism (over 60%) proves the intensity of the speciation processes within the Zerconidae family on the Korean Peninsula.     

Genetic assessment of <Emphasis Type="Italic">Abies koreana</Emphasis> (Pinaceae), the endangered Korean fir,and conservation implications

To establish a management plan for endangered and rare species, genetic assessment must first be conducted. The genetic characteristics of plant species are affected by demographic history, reproductive strategy, and distributional range as well as anthropological effects. Abies koreana E. H. Wilson (Pinaceae), Korean fir, is endemic to Korea and found only in sub-alpine areas of the southern Korean Peninsula and Jejudo Island. This species has been designated as critically endangered by the International Union for Conservation of Nature due to a continuous decline in its range and population fragmentation. We genotyped 176 individuals from seven natural populations and two afforested populations on the Korean Peninsula using 19 microsatellite loci. STRUCTURE analysis revealed two genetic clusters in natural populations (F _st  = 0.040 and R _st  = 0.040) despite low differentiation. We did not detect a significant reduction in genetic diversity or the signature of a genetic bottleneck despite population fragmentation and small population size. We deduced that this species exhibits a metapopulation structure, with the population on Jirisan Mountain acting as a source of genetic diversity for other local small populations on the Korean Peninsula, through contemporary asymmetric gene flow. However, the majority of afforested individuals on the Korean Peninsula originated from a different gene cluster. Thus, we recommend a conservation strategy that maintains two genetically unique clusters.     

Anatomy of the antennal dorsal organ in female of Neodryinus typhlocybae (Hymenoptera: Dryinidae): A peculiar sensory structure possibly involved in perception of host vibration

Neodryinus typhlocybae (Hymenoptera: Dryinidae) is a natural enemy of the planthopper Metcalfa pruinosa, which was introduced from North America into Europe and has become established in various regions as a pest species. Vibrational signals play a crucial role in the communication of M. pruinosa, which appears to be exploited by N. typhlocybae. Scanning and transmission electron microscopy have shown that the antennae of N. typhlocybae females have peculiar and complex sensory structures: deep longitudinal grooves that house long sensilla trichodea, termed here “Antennal Dorsal Organs.” Such structures were not present on male antennae. These sensilla extend for the length of the grooves, without contact with the groove cuticle. Their hair shaft is empty and aporous, and inserted into a specialized socket, underneath which there is a cuticular ampulla‐like chamber. Each sensillum is associated with two sensory neurons: one terminates at the proximal end of the dendritic sheath; the other continues into the sensillum sinus and is enclosed in the dendritic sheath. This second sensory neuron then enters the ampulla‐like chamber through the circular opening, and then terminates with a conspicuous tubular body at the shaft base. The possible involvement of this peculiar structure in the context of host recognition mechanism is discussed. J. Morphol. 277:128–137, 2016. © 2015 Wiley Periodicals, Inc.     

A new record of the Naratettix rubrovittatus () (Hemiptera: Cicadellidae) from Korea

Naratettix rubrovittatus (Matsumura, 1920) is recognized for the first time in Korea on Rhododendron indicum, Rhododendron mucronulatum, Rhododendron yedoense var. poukhanense, and Viburnum opulus var. calvescens. The male specimen is redescribed, illustrated, and measured. To date, three Naratettix species have been recorded on the Korean Peninsula. A key to species of the genus Naratettix from the Korean Peninsula is provided.     

One new record of the genus Aulacorthum (Hemiptera: Sternorrhyncha: Aphididae) from Korea

Aulacorthum vandenboschi was recognized for the first time in Korea on Cirsium japonicum var. ussuriense in Jeju-do. The apterous viviparous female is redescribed, illustrated, and measured. To date, 15 species of the genus Aulacorthum have been recorded on the Korean Peninsula. A key to species of the genus Aulacorthum from the Korean Peninsula is provided.