Isolation and characterization of eight microsatellite loci from <Emphasis Type="Italic">Lycorma delicatula</Emphasis> (White) (Hemiptera: Fulgoridae) for population genetic analysis in Korea

Lycorma delicatula (White) is native to China but is becoming an important insect pest in Korea. Polymorphic DNA markers like microsatellites are widely used for characterizing dispersal patterns and capacity of invasive insect pests which can contribute to designing effective management of the species. To facilitate such population genetic studies of L. delicatula in Korea, we isolated and characterized eight microsatellite loci for L. delicatula using a hybridization-biotin enrichment method. We further used these novel microsatellite loci to determine population genetic parameters for 33 L. delicatula specimens collected from Cheonan, South Korea where outbreaks of this species were first reported in Korea. The number of alleles per locus ranged from three to ten, with an average of 6.25. The mean expected (H _E) and observed heterozygosities (H _O) were 0.575 and 0.626, respectively. The eight loci showed no deviation from Hardy–Weinberg equilibrium according to the adjusted significance threshold (P = 0.00625), and there was no linkage disequilibrium between each pair of these eight markers. Bayesian cluster analysis using the program structure revealed no evidence of genetic structuring in L. delicatula samples from Cheonan. These new microsatellite markers will be widely applicable to future ecological genetic studies of L. delicatula, including assessment of the level of gene flow and genetic connectivity among populations that are necessary for effective management and monitoring of the species.     

A review of biology and management of Lycorma delicatula (Hemiptera: Fulgoridae), an emerging global invasive species

Lycorma delicatula (White) (Hemiptera: Fulgoridae) is an invasive species inflicting damage to agricultural and forest systems. This species is native to China, Taiwan, and Vietnam. Lycorma delicatula was accidentally introduced into South Korea and Japan, with outbreaks reported since mid-2000s. More recently, this pest was found in Pennsylvania, USA, and its geographical distribution has steadily expanded since the introduction with increasing population densities in the USA. This species is univoltine in invaded regions and overwinters in the egg stage. Lycorma delicatula is known to attack at least 65 host plants, and so-called preferred host plants such as Ailanthus altissima and Vitis vinifera are present in all invaded countries. This species threatens grape industries in China, South Korea, and USA, with the impact on other crops still unknown, at least in the USA. For monitoring this pest, light or pheromone traps have not yet been developed, but using sticky tree bands may serve as a promising tool. To date, visual survey is the primary means for detecting L. delicatula and studying their population dynamics. To control L. delicatula, several studies evaluated conventional insecticides. Under field conditions, chlorpyrifos, dinotefuran, etofenprox, and etofenprox + diazinon yielded high levels of mortality against this pest. In general, environmentally-friendly materials, registered as Environmentally-Friendly Agricultural Material in South Korea, resulted in lower insecticidal efficacy against L. delicatula compared with conventional chemicals. Natural enemies of L. delicatula such as egg parasitoids in the genus Anastatus and Dryinus are promising candidates as a long-term solution for management.     

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.     

Spectral preferences of Lycorma delicatula (Hemiptera: Fulgoridae)

Many insects prefer lights with certain spectral properties, and such preferences may be associated with behavioral contexts such as mating, host finding or dispersal. Lycorma delicatula (Hemiptera: Fulgoridae) is a newly invasive species in Korea and is spreading rapidly. It is diurnal and may rely on visual cues for orientation. We conducted a series of three phototaxis experiments to understand spectral preferences in L. delicatula: light/dark choice, UV/white light choice, and color preference experiments. Nymphs of the third and final stages as well as adults were used for these experiments. In the light/dark choice, the orientation of L. delicatula was bimodal between the white light and darkness, for all stages and both sexes. In a choice of UV (395–410 nm wavelengths) vs. white light, L. delicatula of both sexes and all stages preferred the UV light. In the color preference experiment where insects had a choice of four colors in a circular arena, L. delicatula stayed significantly longer in the blue light than in white, yellow or green lights. Overall, nymphs and adults of L. delicatula oriented toward lights with shorter wavelengths, and this orientation was consistent throughout all stages, regardless of sex. It is necessary to investigate the behavioral contexts under which L. delicatula prefers the UV and blue lights.     

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.     

Molecular comparison of Lycorma delicatula (Hemiptera: Fulgoridae) isolates in Korea,China, and Japan

Lycorma delicatula (White) was recently introduced in Korea, acting as a grape insect pest. Since the introduction of this invasive insect, it initially spread rapidly throughout central and southern Korea, and is now distributed throughout the mainland. Here we developed new mitochondrial markers from NADH dehydrogenase subunit 2 and NADH dehydrogenase subunit 6 regions, and analyzed the regional isolates of L. delicatula collected from original locations in China, as well as invasive locations in Korea and Japan. All Korean and Japanese isolates were found to be genetically identical to those from Beijing, Tianjin, Qingdao, and Shanghai, China. Further isolates, from Zhejiang province, China, had two additional haplotypes.     

Cyclic Behavior of <Emphasis Type="Italic">Lycorma delicatula</Emphasis> (Insecta: Hemiptera: Fulgoridae) on Host Plants

Lycorma delicatula is an invasive insect species in Korea, and its populations are increasing. It sucks plant sap and can cause withering of the whole trees. To prevent damage to fruit trees, it is important to determine the behavioral characteristics of L. delicatula. We monitored migration patterns and host plant preferences, from the egg mass on the tree bark, through egg hatching to egg laying. Most eggs hatched between 0500 h and 0800 h. The nymphs ascended trees until they reached the leaves. They often fell to the ground because of physical factors, such as wind, but then ascended the trees again on the upper sides of the branches. Their falling-ascending cycle became longer as their arolia developed. The host preference changed over the course of its growth, from a broad range of plant species at the early nymph stage to a few plant species, such as Ailanthus altissima at the adult stage. We summarize the life cycle behavior as follows. Initially, there is a short-term cycle of falling and ascending, which becomes longer as the nymphs age. The other cycle is a yearly cycle of host plant preferences, with broad host preference in the early nymph stage and a narrower host preference in the adult stage. Knowledge of this cyclic behavior can be used to prevent the rapid expansion of L. delicatula populations in orchards.     

Moving eDNA surveys onto land: Strategies for active eDNA aggregation to detect invasive forest insects

The use of environmental DNA (eDNA) surveys to monitor terrestrial species has been relatively limited, with successful implementations still confined to sampling DNA from natural or artificial water bodies and soil. Sampling water for eDNA depends on proximity to or availability of water, whereas eDNA from soil is limited in its spatial scale due to the large quantities necessary for processing and difficulty in doing so. These challenges limit the widespread use of eDNA in several systems, such as surveying forests for invasive insects. We developed two new eDNA aggregation approaches that overcome the challenges of above‐ground terrestrial sampling and eliminate the dependency on creating or utilizing pre‐existing water bodies to conduct eDNA sampling. The first, “spray aggregation,” uses spray action to remove eDNA from surface substrates and was developed for shrubs and other understorey vegetation, while the second, “tree rolling,” uses physical transfer via a roller to remove eDNA from the surface of tree trunks and large branches. We tested these approaches by surveying for spotted lanternfly, Lycorma delicatula, a recent invasive pest of northeastern USA that is considered a significant ecological and economic threat to forests and agriculture. We found that our terrestrial eDNA surveys matched visual surveys, but also detected L. delicatula presence ahead of visual surveys, indicating increased sensitivity of terrestrial eDNA surveys over currently used methodology. The terrestrial eDNA approaches we describe can be adapted for use in surveying a variety of forest insects and represent a novel strategy for surveying terrestrial biodiversity.     

Orientation of flight for physically disturbed spotted lanternflies,Lycorma delicatula, (Hemiptera,Fulgoridae)

The spotted lanternfly, Lycorma delicatula, (Hemiptera, Fulgoridae) is an invasive pest to Korea and the United States, originating from China or Southeast Asia. Immature L. delicatula feed on a wide range of plants, but the adults are more host-selective, often preferring the tree of heaven, Ailanthus altissima. We performed field studies to evaluate adult movement in relation to A. altissima after disturbance. The Ailanthus trees were in a mixed suburban forested situated at the southern border of an open grassy field. Female adult lanternflies were manually disturbed from feeding on the trunks of large A. altissima trees. In the first experiment, insects were disturbed directly from the tree using a ballpoint pen to simulate a predatory attack. These insects usually flew initially southward away from the tree line toward a sunlit field, but turned northward back toward the tree line. In the second experiment, to simulate an initially successful predatory attack, they were manually taken from trees, and allowed to escape. The females immediately opened their wings in an apparent aposematic display. They then either immediately flew toward the sunlit open field, or remained with their wings splayed open for a prolonged period.     

Skewed adult sex ratios observed early in the North American invasion of Lycorma delicatula (Hemiptera: Fulgoridae)

The spotted lanternfly, Lycorma delicatula, (Hemiptera Fulgoridae) is an invasive pest in Korea and the United States, but originating from mainland Asia. Both tree of heaven, Ailanthus altissima, and grapevine (Vitis) are known to be preferred hosts of adults. However, much is unknown about its adult behavior with respect to mating or dispersal patterns. In 2015, just after it was discovered in the United States, we performed an observational study to elucidate movement by adults in relation to host plants. We noted weekly changes in their presence and sex ratio on host plants at four sites in a quarantine zone in Berks County, PA. Just after adult emergence, the trunks of large (>15 cm diameter at breast height) A. altissima hosted nearly exclusively females, while smaller A. altissima, Vitis, Salix nigra, and other less commonly inhabited tree species, had high proportions of males. By October 1st, greater proportions of males were observed on the trunks of the larger A. altissima trees. Pairings of males and females were observed most frequently on the smaller A. altissima and Vitis. At the very end of the season, a large mixed-sex population was found on Salix nigra, which was the only deciduous tree species in the area still bearing green leaves.     

Geographical distribution of bat flies (Diptera: Nycteribiidae and Streblidae), including two new records,Nycteribia allotopa and N. formosana,collected from bats (Chiroptera: Rhinolophidae and Vespertilionidae) in the Republic of Korea

As part of the 65^th Medical Brigade, U.S. Army, arthropod‐borne disease surveillance program and in collaboration with the Korea National Institute of Biological Resources (NIBR), bats were captured from caves and abandoned mines in the Republic of Korea. A total of 39 adult bat flies including five species of Nycteribiidae [Penicillidia jenynsii, Nycteribia parvula, N. formosana, N. allotopa mikado, and an unidentified species of Nycteribia (N. cf. formosana)], and one species of Streblidae, Brachytarsina kanoi, were collected from bats belonging to two families, Rhinolophidae and Vespertilionidae. This is the first report of N. allotopa mikado and N. formosana from the Republic of Korea.     

Impact of minimum winter temperature on Lycorma delicatula (Hemiptera: Fulgoridae) egg mortality

Climate change, especially winter temperature increase, may be a factor the recent occurrence and range expansion of Lycorma delicatula (White) in South Korea. Egg mortality increased as minimum winter temperature decreased. Egg mortality was highest (93.32%) in Chuncheon, where the mean minimum winter temperature was lowest. The temperature that induced complete mortality of fulgorid eggs was estimated to be ?3.44 °C on the basis of mean daily temperatures from December 2009 to February 2010 and ?12.72 °C on the basis of mean daily minimum temperatures in January 2010. Mean winter temperatures during the last 30 years have gradually increased to above the temperature for 100% mortality of the fulgorid egg. This suggests that the winter temperature increase may contribute to the successful settlement of the fulgorid in South Korea and its subsequent outbreak and expansion.     

Rediscovery of Ardisia gigantifolia and the reinstatement of A. kteniophylla (Primulaceae) – finding the correct name for the Chinese medicinal plant ’Zou Ma Tai’

With the rediscovery of Ardisia gigantifolia, the long‐standing confusion with A. pseudoverticillata, A. kteniophylla and A. dasyrhizomatica owing to the misinterpretation of the type of A. gigantifolia is clarified. It is shown that A. dasyrhizomatica is a later synonym of A. gigantifolia, while A. kteniophylla and A. pseudoverticillata are conspecific and represent another species for which A. kteniophylla is the earliest name available. Ardisia kteniophylla is the correct scientific name for the Chinese medicinal plant ’Zou Ma Tai’.     

Morphology‐based phylogenetic binning to assess a taxonomic challenge: a case study in Graphidaceae (Ascomycota) requires a new generic name for the widespread Leptotrema wightii

Molecular phylogenetic analysis presents two challenges when it is transformed into formal classifications: the taxonomic challenge (whether and how to distinguish monophyletic sister clades or how to deal with paraphyletic grades) and the nomenclatural challenge (naming clades, i.e. placing name‐giving types accurately on a tree). One approach to the latter is morphology‐based phylogenetic binning, which places specimens based on phenotypic features on a molecular tree and assigns uncertainty values to alternative placement options. Here, we use the example of the lichenized fungal genus Leptotrema to demonstrate how morphology‐based phylogenetic binning can help to clarify taxonomic and nomenclatural issues when naming phylogenetically defined entities. Leptotrema is known for a common and widespread species, L. wightii, and phylogenetic analyses have been based exclusively on this species, including the recognition of a separate tribe, Leptotremateae. However, the genus name Leptotrema and the tribal name Leptotremateae are based on the name L. zollingeri, which was initially considered to be a synonym of L. wightii, but has recently been shown to represent a distinct species. As L. zollingeri differs considerably in phenotypic features from L. wightii, it can be questioned whether the two are at all related or whether L. zollingeri is actually closer to the genera Myriotrema and Ocellularia in tribe Ocellularieae. The solution to this problem is not trivial, as it affects the correct use of the names Leptotrema and Leptotremateae. Morphology‐based phylogenetic binning indeed demonstrated that L. zollingeri clusters with the Myriotrema album group in tribe Ocellularieae with high support. Hence, in contrast with current use, the name Leptotrema becomes available for the M. album group and Leptotremateae becomes a synonym of Ocellularieae. As a consequence, the new names Sanguinotrema and Sanguinotremateae are introduced to accommodate L. wightii and the tribe including this species and the genus Reimnitzia. Although the studied case is specific to lichen fungi, the approach can be used in a much broader context with any kind of taxon or organism. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 436–443.     

DEVELOPMENTAL STUDY OF BRANCHED RHIZOPHORES IN THREE SELAGINELLA SPECIES

A morphogenetic investigation was made of the rhizophore of three large-sized tropical Selaginella species. The rhizophores of Selaginella delicatula, S. caudata, and S. plana arise exogenously at the points of branching of the main stems. In S. delicatula they are initiated at the junction of the second youngest branching. The rhizophore apical meristem has a tetrahedral apical cell and is capless. The rhizophores are usually three or four times dichotomously branched in S. delicatula and S. plana and four or five times in S. caudata. In S. delicatula, dichotomous branching of the rhizophore involves formation of two new apical cells subsequent to loss of an original apical cell. A pair of roots is formed endogenously from inner cells below the dermal layer at the apex of ultimate rhizophore branches. The finding that the rhizophore is an autonomously branched, leafless, and capless axis leads us to argue that Selaginella rhizophores, like lepidodendrid rhizomorphs, are fundamental axial organs that coordinate with the stem and root.     

Revision of the family Sphecidae (Hymenoptera: Apoidea) in South Korea

Korean species of the family Sphecidae are reviewed, and seventeen species in nine genera belonging to three subfamilies are confirmed. Among them, Ammophila campestris and Sceliphron caementarium are new to South Korea. Korean occurrence of Sceliphron madraspatanum is confirmed with an actual Korean specimen. This paper provides the first determination keys to higher taxa and species occurring in South Korea. Taxonomic information of each species including original publication of valid name and justifiable references for Korean occurrence is provided. Digital images and line drawings for selective characteristics are also available.     

A Molecular Phylogeny of Hemiptera Inferred from Mitochondrial Genome Sequences

Classically, Hemiptera is comprised of two suborders: Homoptera and Heteroptera. Homoptera includes Cicadomorpha, Fulgoromorpha and Sternorrhyncha. However, according to previous molecular phylogenetic studies based on 18S rDNA, Fulgoromorpha has a closer relationship to Heteroptera than to other hemipterans, leaving Homoptera as paraphyletic. Therefore, the position of Fulgoromorpha is important for studying phylogenetic structure of Hemiptera. We inferred the evolutionary affiliations of twenty-five superfamilies of Hemiptera using mitochondrial protein-coding genes and rRNAs. We sequenced three mitogenomes, from Pyrops candelaria, Lycorma delicatula and Ricania marginalis, representing two additional families in Fulgoromorpha. Pyrops and Lycorma are representatives of an additional major family Fulgoridae in Fulgoromorpha, whereas Ricania is a second representative of the highly derived clade Ricaniidae. The organization and size of these mitogenomes are similar to those of the sequenced fulgoroid species. Our consensus phylogeny of Hemiptera largely supported the relationships (((Fulgoromorpha,Sternorrhyncha),Cicadomorpha),Heteroptera), and thus supported the classic phylogeny of Hemiptera. Selection of optimal evolutionary models (exclusion and inclusion of two rRNA genes or of third codon positions of protein-coding genes) demonstrated that rapidly evolving and saturated sites should be removed from the analyses.     

Studies in African Cyperaceae 37: Scleria aurantiaca sp. nov. from Gabon

Scleria aurantiaca Lye is described from eastern Gabon. It is superficially similar to S. nyassensis, but probably more closely related to S. adpresso‐hirta, which has a more similar nutlet and hypogynium. The scientific name is derived from the bright orange ring of the hypogynium. The fruit of S. aurantiaca has both micro‐ and nano‐papillae, which are described here for the first time. Also discussed is the water transport through surface hairs to the embryo in the fruit. Due to the restricted range of this species, it must be considered as ‘Endangered’ (EN).     

Larval descriptions of five Byctiscus species (Coleoptera: Rhynchitidae) in Korea

The genus Byctiscus reported in Korea in five of nine species (B. lacunipennis, B. rugosus, B. puberulus puberulus, B. princeps and B. coerulans) is described for the last instar larvae. In addition, the cradle structure and host‐plants are provided.