Genetic diversity of the brown marmorated stink bug <Emphasis Type="Italic">Halyomorpha halys</Emphasis> in the invaded territories of Europe and its patterns of diffusion in Italy

Halyomorpha halys is an invasive stink bug pest originating from East Asia. In Europe, it was first detected in Switzerland in 2004. It is now present in thirteen countries, and seems to be spreading throughout the continent. In Italy, where it has been recorded since 2012, other than being an urban nuisance, it is causing severe damage in commercial fruit orchards. An integrated approach, using current and previous observational data in space and time and molecular information, was used to identify the genetic diversity of this pest in Europe, its invasion history, and the potential pathways of entry and diffusion. The analysis of 1175 bp of mitochondrial DNA cytochrome c oxidase I and II genes (cox1, cox2) led to the identification of twenty previously unknown haplotypes. The European distribution of H. halys is the result of multiple invasions that are still in progress, and, in some cases, it was possible to identify the specific Asian areas of origin. Moreover, secondary invasions could have occurred among European countries by a bridgehead effect. In Italy, the data were more clearly related to their temporal occurrence, allowing for a clearer reading of the patterns of invasion and dispersion. After having successfully established in localized areas, H. halys further expanded its range by an active dispersion process and/or by jump dispersal events due to passive transport. The multiple introductions from different areas of the native range together with the different patterns of diffusion of H. halys, may hamper the pest management strategies for its containment.     

Predation of <Emphasis Type="Italic">Halyomorpha halys</Emphasis> (Hemiptera: Pentatomidae) from Web-Building Spiders Associated with Anthropogenic Dwellings

The brown marmorated stink bug, or Halyomorpha halys, is an invasive pest in North America and Europe that causes severe agricultural damage and nuisance problems for homeowners; and it is originally from China, Taiwan, and the Republic of Korea. While the natural enemy community of H. halys has been evaluated in several agroecosystems, it has not been examined where H. halys overwinters in anthropogenic structures. The aims of the current study were to evaluate 1) whether spider webs commonly found in the home and yard can successfully ensnare H. halys, 2) whether entanglement resulted in consumption by spiders inhabiting the webs, and 3) how frequently H. halys becomes entangled in webs under ambient conditions. To accomplish this, adult H. halys were introduced into webs in and near anthropogenic structures in West Virginia and Maryland, United States, and the behavior of spiders was observed for 5-min intervals at 0, 1, 2, and 24 h after introduction. In addition, a survey of webs was performed to determine the frequency with which spiders naturally capture H. halys inside buildings and in the landscape. Overall, the study found seven spider families in anthropogenic structures. Adult H. halys that were introduced into the webs of Theridiidae, Pholcidae, or Agelenidae had a greater than 50% chance of being ensnared and consumed. Adult H. halys were found naturally most often in webs of Theridiidae. Webs with a funnel or cob web architecture had the greatest probability of ensnaring H. halys, while those with orb structures resulted in the fewest caught. In the wild, 13–20% of spider webs contained dead H. halys. Our results suggest that spiders may be an important contributing factor for mortality of H. halys at overwintering sites, and spiders in or outside homes may help reduce nuisance problems caused by H. halys.     

Review of the occurrence of <Emphasis Type="Italic">Halyomorpha halys</Emphasis> (Hemiptera: Heteroptera: Pentatomidae) in Italy,with an update of its European and World distribution

The regions in which Halyomorpha halys (Stål, 1855) occurs in Italy are summarized and ordered chronologically according to the date of the first record. Details and verification about the first Italian record from Liguria (often overlooked), that dates back to 2007 the introduction of H. halys in this country, are also given. First exact localities are listed for the Italian regions of Tuscany, Marche, Latium, Abruzzo, Campania, and Apulia, in which only general occurrence was known so far. We included also additional records from Trentino-Alto Adige, where the presence of H. halys has only recently been reported through a few data. Furthermore first records for Sweden, Central Macedonia (Greece), and Manitoba (Canada) are reported, as well as first intercepted specimens from Iceland.     

Invasive brown marmorated stink bug <Emphasis Type="Italic">Halyomorpha halys</Emphasis> (Stål) (Heteroptera: Pentatomidae) in Russia,Abkhazia, and Serbia: history of invasion,range expansion,early stages of establishment,and first records of damage to local crops

Halyomorpha halys is native to Asia, but was recently accidentally introduced to North America and Europe. This species was recorded for the first time in Russia in 2013 (2014) and in both Serbia and Abkhazia in 2015. In 2015–2017, we conducted surveys in Russia, Serbia, and Abkhazia and found that H. halys had not only successfully survived its first winters in new regions, but also started to establish locally, spread, increase populations, and damage local crops. In Sochi (Russia) and Abkhazia, H. halys was recorded to feed on 32 species and cultivars from 16 plants families. In Serbia, it has so far been observed on only four species from four plant families. H. halys is already widely spread in Abkhazia and causes serious damage to many crops: in 2016, the yield of peach, mandarin orange, persimmon and other crops fell down by 13.2–87.4% if compared to the long-term means of yield. The losses are likely to be mostly caused by feeding of H. halys. We conclude that H. halys was accidentally introduced to Russia in 2012–2013 most likely with woody plants imported to Sochi from Italy or Greece for urban landscaping before the 2014 Winter Olympics. The species established in Sochi and quickly moved to Krasnodar (Russia), Abkhazia, and beyond—to Georgia. Populations in Serbia most likely arrived in 2015 traveling on railway cars or in commodities on the Bucharest–Belgrade railway line. We discuss phenology and prospects of further expansion of H. halys in Europe.     

Modelling the potential geographic distribution of <Emphasis Type="Italic">Trissolcus japonicus</Emphasis>: a biological control agent of the brown marmorated stink bug, <Emphasis Type="Italic">Halyomorpha halys</Emphasis>

Trissolcus japonicus Ashmead (Hymenoptera: Scelionidae) is an endoparasitoid of the eggs of the brown marmorated stink bug, Halyomorpha halys Stål (Hemiptera: Pentatomidae), a major agricultural pest native to China, Japan, South Korea and Taiwan. We used CLIMEX to estimate the potential global distribution of T. japonicus with particular reference to New Zealand. In its native range the model predicts the presence, or a potential expansion, of T. japonicus into most of humid-subtropical and humid-continental areas. Globally, the model projects that many temperate, Mediterranean and subtropical areas could suit the establishment of T. japonicus. In New Zealand, the north appears moderately to highly suitable for T. japonicus, while southern regions are mostly marginal. The risk posed by T. japonicus to non-target species in New Zealand is predicted to vary between different non-targets. CLIMEX projections of the potential distribution of T. japonicus provide guidance for release sites of this parasitoid if approved for importation and release in New Zealand.     

First records of the brown marmorated stink bug <Emphasis Type="Italic">Halyomorpha halys</Emphasis> (Stål, 1855) (Heteroptera,Pentatomidae) in Russia,Abkhazia, and Georgia

The brown marmorated stink bug Halyomorpha halys (Stål, 1855) is recorded for the first time from Russia (Khosta and Adler districts of the city of Sochi), Abkhazia (Pitsunda), and Georgia (Khobi Municipality). Outbreaks of H. halys populations were detected in all of these regions in October 2016. More than a hundred specimens congregated for overwintering were collected in Sochi during two days. According to the personal communications of the local people, the species appeared in Sochi not later than in 2013, and in Georgia not later than in 2015. It must have been brought to Sochi from Europe (from Italy) with planting material of ornamental plants for landscaping the XXII Olympic Winter Games. Thereafter it spread to Abkhazia and Georgia. Additional steps to identification of H. halys are provided for the keys to the true bugs of the European part of the former USSR (Kerzhner and Jaczewski, 1964).     

Control effect on the brown-marmorated stink bug, <Emphasis Type="Italic">Halyomorpha halys</Emphasis> (Hemiptera: Pentatomidae), by combined spraying of pyrethroid and neonicotinoid insecticides in apple orchards in northern Japan

Control of Halyomorpha halys (Stål) adults by combined spraying of pyrethroid and neonicotinoid insecticides has been investigated in apple orchards in northern Japan in 2008 and 2009. Insecticides were selected to control not only H. halys but also the common lepidopteran pests Carposina sasakii Matsumura, Adoxophyes orana fasciata Walsingham, and Phyllonorycter ringoniella Matsumura. Dinotefuran was sprayed in mid-May and early July, fenpropathrin in late June, and bifenthrin in late July. H. halys adults were released into apple trees at 3–5-day intervals from mid-May to mid-August. In this spraying program, survival of the adults increased 6 days after spraying, but relatively few injury marks were noted throughout the growing season. On the other hand, in the conventional spraying program, the survival ratio of the adults increased rapidly 3 days after spraying—especially after the organophosphate applied in mid-May, late June, and late July—and increases in the number of injury marks synchronized well with survival in both years. This spray program reduced the number of fruit injury marks to one-fifth that of the conventional spraying program and one-tenth that of fungicides alone.     

Complete mitochondrial genome of brown marmorated stink bug <Emphasis Type="Italic">Halyomorpha halys</Emphasis> (Hemiptera: Pentatomidae), and phylogenetic relationships of hemipteran suborders

The newly sequenced complete mitochondrial genome of the brown marmorated stink bug, Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), is a circular molecule of 16,518 bp with a total A+T content of 76.4% and two extensive repeat regions in A+T rich region. Nucleotide composition and codon usage of H. halys are about average when compared with values observed in 19 other published hemipteran mitochondrial genomes. Phylogenetic analyses using these 20 hemipteran mitochondrial genomes support the currently accepted hypothesis that suborders Heteroptera and Auchenorrhyncha form a monophyletic group. The mitochondrial gene arrangements of the 20 genomes are also consistent with our results.     

<Emphasis Type="Italic">Hexamermis popilliae</Emphasis> n. sp. (Nematoda: Mermithidae) parasitizing the Japanese beetle <Emphasis Type="Italic">Popillia japonica</Emphasis> Newman (Coleoptera: Scarabaeidae) in Italy

A new species of mermithid nematode, Hexamermis popilliae n. sp. (Nematoda: Mermithidae) is described from the Japanese beetle Popillia japonica Newman in Italy, an area of new introduction for this invasive pest. The combination of the following characters separates H. popilliae from other members of the genus Hexamermis Steiner, 1924: adult head obtuse; amphidial pouches slightly posterior to lateral head papillae in female but adjacent to lateral head papillae in males; amphidial openings large, well developed; amphidial pouches elliptical in females and oblong in males; cuticular vulvar cone well developed, vulvar lips greatly reduced or lacking, vagina curved at tip where meeting uteri, without reverse bend (not S-shaped), spicules slightly curved, with a slight bend in the basal portion, approximately equal to body width at cloaca. This is the first record of a species of Hexamermis parasitizing the Japanese beetle Popillia japonica. The only previous mention of mermithid nematodes from P. japonica was an undescribed species of Psammomermis in North America. Hexamermis popilliae will be evaluated as a potential biological control agent in an integrated control program of the Japanese beetle in Italy.     

Reaping benefits from an invasive species: role of <Emphasis Type="Italic">Harmonia axyridis</Emphasis> in natural biological control of <Emphasis Type="Italic">Aphis glycines</Emphasis> in North America

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is an invasive species present in numerous agroecosystems in North America. Despite adverse impacts as a threat to native biodiversity, a nuisance household invader and a pest in fruit production, H. axyridis also plays a beneficial role as a major component of assemblages of generalist predators in several agricultural crops. Here, we review the role of H. axyridis as a natural enemy of Aphis glycines Matsumura (Hemiptera: Aphididae), an invasive pest of soybean, Glycine max (L.) Merrill (Fabales: Fabaceae), in North America. Harmonia axyridis is often the most abundant predator species attacking A. glycines in soybean agroecosystems. This predator has the potential to both prevent and suppress A. glycines outbreaks. Further studies are needed to fully understand and utilize the potential of H. axyridis as a natural enemy in the management of A. glycines and other agricultural pests in agroecosystems worldwide.     

A new invasive species of buprestid beetles in the Russian fauna: <Emphasis Type="Italic">Lamprodila</Emphasis> (<Emphasis Type="Italic">Palmar</Emphasis>) <Emphasis Type="Italic">festiva</Emphasis> (L.) (Coleoptera,Buprestidae), a pest of Cupressaceae

The buprestid beetle Lamprodila (Palmar) festiva (L.) (Coleoptera, Buprestidae, Poecilonoini), an invasive pest of cultivated Cupressaceae such as Thuja, Chamaecyparis, Cupressus, and Juniperus in the Greater Sochi area (the Russian Black Sea coast), is recorded for the first time for Russia and the former USSR. Data on the systematic position, diagnostic characters, distribution, host plants, biology of the new pest, and its damage to urban plantations are discussed. Recent records of this species in the new areas of Central and Eastern Europe as well as on the Black Sea coast of the Caucasus suggest expansion of its range during the recent decades.     

<Emphasis Type="Italic">Spiroplasma</Emphasis> dominates the microbiome of khapra beetle: comparison with a congener,effects of life stage and temperature

Khapra beetle, Trogoderma granarium (Coleoptera: Dermestidae), is among the world’s most invasive and destructive pests of stored agricultural products. Its pest status is enhanced by the ability of the larvae to undergo diapause, which increases their tolerance to adverse conditions including insecticides and extreme temperatures. The ability of insects to tolerate extreme conditions can be influenced by their associated bacterial community (the microbiome). Understanding this relationship may lead to improved methods of pest control, but the microbiome of T. granarium is unknown. Here we use next-generation sequencing to address three main questions: 1) How similar are the microbiomes of the closely-related species T. granarium and T. variabile? 2) How does the microbiome change across life stage and physiological state? 3) How is the microbiome of adult T. granarium affected by extreme temperatures? Our results show that the core microbiomes of T. granarium and T. variabile are similar in composition. However, adults of former species have a microbiome dominated by Spiroplasma bacteria (99% of amplified sequences), whereas Spiroplasma in the latter species is almost absent (< 2%). The microbiome of T. granarium differs across life stage (feeding vs non-feeding life stages); its presence in eggs confirms the vertical transmission of Spiroplasma. High temperatures significantly reduced the relative abundance of Spiroplasma, but an effect of low temperatures on the microbiome of T. granarium was not detected. Given its dominance in a key pest species, further study of the interaction between Spiroplasma and its T. granarium host is warranted.     

Identifying the ecological and societal consequences of a decline in <Emphasis Type="Italic">Buxus</Emphasis> forests in Europe and the Caucasus

The potential impact of new invasive tree pests and diseases is usually quantified in economic terms. The ecological and social impacts are less often assessed. Using a comprehensive literature review we assess the potential ecological and social impact of two non-native invasive species (the box tree moth, Cydalima perspectalis and the fungus Calonectria pseudonaviculata) that threaten the survival of box tree, Buxus spp. in forests in Europe and the Caucasus. A total of 132 fungi, 12 chromista (algae), 98 invertebrate and 44 lichens were found to use Buxus spp. Of these, 43 fungi, 3 chromista and 18 invertebrate species have only been recorded on Buxus spp., suggesting that these species are obligate on Buxus spp. and are most at risk from in the loss of Buxus spp. due to these invasive pest and disease species. Buxus spp. was shown to be important for soil stability and water quality but there was no information on other ecosystem functions provided by Buxus spp. Buxus was found to be of considerable historical cultural importance but there was very limited information on current social values and uses. Buxus trees, wood and leaves are associated with different folklore and sacred rites which are still particularly important in the Caucasus. While we could not find any assessment of the economic value of Buxus forests the biodiversity, cultural and social values of Buxus identified here indicate that its loss could have major indirect and non-market economic effects. This work highlights the importance of studying the ecological and societal implications of biological invasions.     

The seeds of success: release from fungal attack on seeds may influence the invasiveness of alien <Emphasis Type="Italic">Impatiens</Emphasis>

Although closely related, Impatiens glandulifera and Impatiens balfourii differ in their invasiveness in Europe; only the former is highly invasive there. Following the assumptions of the enemy release hypothesis (ERH), we tested whether these differences may be explained by the levels of seed infestation by pathogenic fungi. Using seeds collected along the Swiss-Italian border, we recorded four true pathogens of seeds: Fusarium culmorum, F. oxysporum, F. sporotrichoides, and Giberella avenacea. In Italy the seeds of I. balfourii were infected by fungal pathogens more often than those of I. glandulifera, while in Switzerland both species were under the same level of pressure. However, the overall differences in pathogen abundance were consistent with the ERH: seeds of the more invasive species were attacked less. This could be a result of differences between the communities of fungal pathogens attacking the seeds of both species in each country. The number of colonies of secondary pathogens (Cladosporium cladosporioides, Alternaria alternata) correlated negatively with the number of colonies of true pathogens; we suggest that the secondary pathogens may have prevented the occurrence of the true pathogens. The reason for the between-country differences in the fungal pathogen communities is unclear. A possible explanation is that Italy and Switzerland differ in their road and green-area maintenance work schemes, which may have influenced pathogen pressure on seeds. This study is one of the few that offers results indicating that release from enemies may be crucial to the invasion success of plants as early as the seed stage.     

A new <Emphasis Type="Italic">Gephyraulus</Emphasis> species (Diptera: Cecidomyiidae) inducing flower bud galls on the European sea rocket <Emphasis Type="Italic">Cakile maritima</Emphasis> Scop. (Brassicaceae)

The European sea rocket Cakile maritima Scop. (Brassicaceae) is a common herb growing on sandy coastlines worldwide and is considered a useful plant because of its medicinal importance, its edibility, and potential as an oilseed crop. However, C. maritima is an invasive plant over a wide range, e.g., eastern South America, North America, northern Iran, Australia and New Zealand, and has a limited number of associated herbivorous insects. During investigations on gall midges (Diptera: Cecidomyiidae) in Egypt, we found a gall midge inducing flower bud galls on C. maritima and preventing fruit production, which suggested that this gall midge is a potential pest of this plant. In this paper, we describe this gall midge species, Gephyraulus zewaili Elsayed and Tokuda sp. nov., as new to science by comparing its morphology with that of close congeners. Partial sequence data of the mitochondrial DNA cytochrome oxidase subunit I gene are also provided.     

<Emphasis Type="Italic">Scirtothrips dorsalis</Emphasis> (Thysanoptera: Thripidae): a Newly Introduced Polyphagous Pest in Northeastern Brazil

We report for the first time the detection of Scirtothrips dorsalis Hood (Thysanoptera: Thripidae) in Brazil and describe the occurrence of the thrips on leaves of ungrafted dwarf-cashew Anacardium occidentale Linnaeus 1753 (Anacardiaceae), maintained into a greenhouse, in the northeastern state of Ceará. This exotic polyphagous species listed as absent quarantine pest in the country is originated in Asia, but since the beginning of this century, it has readily dispersed despite the strict quarantine regulations currently in several countries. Individuals of S. dorsalis identified in Brazil are similar to specimens found in South Africa rather than Asia by virtue of the absence of conspicuous reticulation on the posterior half of the metanotum and the presence of complete lines of microtrichia restricted to the posterior part of the abdominal sternites. Scirtothrips dorsalis is a particularly invasive pest and its introduction represents a potential threat to various crops in Brazil, especially mango.     

A PCR-based analysis of plant DNA reveals the feeding preferences of <Emphasis Type="Italic">Apolygus lucorum</Emphasis> (Heteroptera: Miridae)

The mirid bug Apolygus lucorum (Meyer-Dür) (Heteroptera: Miridae) is a severe pest of cotton and other crops in China. The feeding preferences of this pest are unclear due to its frequent movement among different host plants and the inconspicuous signs of its feeding. Here, we present results of a field trial that used direct observation of bug densities and a PCR-based molecular detection assay to detect plant DNA in bugs to explore relationships between A. lucorum population abundance and its feeding preference between two host plants, Humulus scandens (Loureiro) Merrill and Medicago sativa L. The field-plot samples showed that A. lucorum adults generally prefer flowering host plants. Its density was significantly higher on flowering H. scandens than on seedlings of M. sativa, and a similarly higher bug density was observed on flowering M. sativa than on seedlings of H. scandens. In the laboratory, we designed two pairs of species-specific primers targeting the trnL-F region for H. scandens and M. sativa, respectively. The detectability of plant DNA generally decreased with time post-feeding, and the half-life of plant DNA detection (DS₅₀) in the gut was estimated as 6.26 h for H. scandens and 3.79 h for M. sativa with significant differences between each other. In mirid bugs exposed to seedlings of H. scandens and flowering M. sativa, the detection rate of M. sativa DNA was significantly higher than that of H. scandens. Meanwhile, in mirid bugs exposed to seedlings of M. sativa and flowering H. scandens, a significantly higher detection rate of H. scandens DNA was found. We developed a useful tool to detect the remaining plant food species specifically from the gut of A. lucorum in the current study. We provided direct evidence of its feeding preference between H. scandens and M. sativa at different growth stages, which strongly supported a positive correlation between population abundance and feeding preference of A. lucorum on different plants under field conditions. The findings provide new insights into the understanding of A. lucorum’s feeding preference, and are helpful for developing the strategies to control this pest.     

<Emphasis Type="Italic">Drosophila</Emphasis> as models to understand the adaptive process during invasion

The last few decades have seen a growing number of species invasions globally, including many insect species. In drosophilids, there are several examples of successful invasions, i.e. Zaprionus indianus and Drosophila subobscura some decades ago, but the most recent and prominent example is the invasion of Europe and North America by the pest species, Drosophila suzukii. During the invasive process, species often encounter diverse environmental conditions that they must respond to, either through rapid genetic adaptive shifts or phenotypic plasticity, or by some combination of both. Consequently, invasive species constitute powerful models for investigating various questions related to the adaptive processes that underpin successful invasions. In this paper, we highlight how Drosophila have been and remain a valuable model group for understanding these underlying adaptive processes, and how they enable insight into key questions in invasion biology, including how quickly adaptive responses can occur when species are faced with new environmental conditions.     

Rearing system for the predatory phytoseiid <Emphasis Type="Italic">Euseius concordis</Emphasis> (Acari: Phytoseiidae)

Several species of predatory mites, especially those of the family Phytoseiidae, are potentially useful for the control of pest mites and insects. Among the phytoseiids, Euseius concordis (Chant), a species commonly found in South America, has been studied for possible use as biological control agent of pest mites. Given that Euseius species are known to feed on plant leaves and on pollen, the objective of this study was to determine plant species and pollen sources suitable to establish a pilot method of production of that species, based on a set of comparisons in the laboratory. Out of four plant species evaluated in the study, higher survivorship of E. concordis females in the absence of supplementary food was observed on leaflets of Canavalia ensiformis L. Out of two pollen types, higher survivorship was obtained on pollen of Typha domingensis L. In a subsequent step, the population of E. concordis increased 19.3× within 21 days when that pollen of T. domingensis was offered to the predator on plantlets of C. ensiformis, at 25.0 ± 1 °C, 70.0 ± 10% RH and 12 h photophase. Future studies may confirm the economic viability of this setup for the mass production of E. concordis.     

<Emphasis Type="Italic">Tamarix</Emphasis> (Tamaricaceae) hybrids: the dominant invasive genotype in southern Africa

Hybridization is regarded as a rapid mechanism for increasing genetic variation that can potentially enhance invasiveness. Tamarix hybrids appear to be the dominant genotypes in their invasions. Exotic Tamarix are declared invasive in South Africa and the exotic T. chinensis and T. ramosissima are known to hybridize between themselves, and with the indigenous T. usneoides. However, until now, it was not known which species or hybrid is the most prevalent in the invasion. With a biocontrol programme being considered as a way of suppressing the alien Tamarix populations, it is important to document the population genetic dynamics of all species in the region. This investigation sought to identify Tamarix species in southern Africa and their hybrids, describe their population structure, and reveal the geographic origin of the invasive species. To achieve this, nuclear Internal Transcribed Spacer (ITS) sequence data and the multilocus Amplified Fragment Length Polymorphisms (AFLPs) markers were used. Phylogenetic analysis and population genetic structure confirmed the presence of three species in South Africa (T. chinensis, T. ramosissima and T. usneoides) with their hybrids. The indigenous T. usneoides is clearly genetically distant from the alien species T. chinensis and T. ramosissima. Interestingly, the Tamarix infestation in South Africa is dominated (64.7 %) by hybrids between T. chinensis and T. ramosissima. The exotic species match their counterparts from their places of origin in Eurasia, as well as those forming part of the invasion in the US.