Specific biochemical marker-based techniques for the identification of damage to Douglas-fir seed resulting from feeding by the western conifer seed bug, Leptoglossus occidentalis Heidemann (Hemiptera: Coreidae)

Specific biochemical marker-based techniques were tested for their ability to distinguish between seeds of Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco, that were filled or unfilled (aborted) at maturity and those that were damaged or emptied by the western conifer seed bug, Leptoglossus occidentalis Heidemann. A polyclonal antibody raised against salivary gland extracts from L. occidentalis successfully identified residual salivary proteins on Western blots containing proteins from Douglas-fir seeds that had sustained various degrees of seed bug feeding damage. In a single blind experiment, the polyclonal antibody correctly identified 100% of undamaged control, 97% of unfilled control (aborted), and 98% of seed bug damaged seeds. Polyclonal antibodies raised against insoluble alfalfa crystalloid storage protein (11S globulin) detected the depletion of 11S globulin and the subsequent appearance of its hydrolyzed fragments in the soluble protein fraction of Douglas-fir seeds that were fed-upon by the seed bug. Feeding by L. occidentalis nymphs caused ca. 98% depletion of insoluble protein, but only ca. 53% reduction in the amount of soluble protein in seeds that appeared empty on radiographs. By comparison, unfilled (aborted) seeds contained significantly less insoluble and soluble protein than empty seeds that were fed-upon by L. occidentalis; moreover, no crystalloid (11S globulin) breakdown products were generated. The biochemical markers described in this study are reliable tools that can be used to identify conifer seeds that have sustained light to severe damage from L. occidentalis feeding.     

Measuring the impact of Leptoglossus occidentalis (Heteroptera: Coreidae) on seed production in lodgepole pine using an antibody-based assay

We measured the impact of Leptoglossus occidentalis on seed production in lodgepole pine, Pinus contorta variety latifolia Engelmann, using an antibody marker developed to detect residual saliva in fed-on seeds. Nymphs, adult females, and adult males were caged on cones during early, mid- and late season cone development. Individual analysis of 12,887 seeds extracted from 365 cones revealed that 37.3% seeds tested positive for seed bug saliva. The antibody assay was 38 times more effective than radiography at detecting seed bug damage. Radiography can detect partially emptied seed but cannot discriminate between aborted seeds and those emptied by seed bugs. The antibody marker was least sensitive in detecting early season damage compared with mid- and late season damage. We hypothesize that residual saliva in seeds fed on early in the season was either absorbed by the damaged seed or degraded over time. Early season feeding resulted in the greatest number of seeds fused to cone scales and the extraction efficiency for cones exposed to feeding during this time was reduced by 64% compared with control cones. Adding fused seeds to antibody-positive seeds raised the proportion of damaged seeds to 48.3%. At all stages of cone development, adult females were the most destructive life stage, damaging up to two seeds per day late in the season. When seed losses were adjusted to damage per degree-day, female damage was greatest early in the season, while males caused the same amount of damage regardless of cone development period. The results of the antibody assay provide baseline data for developing damage prediction formulae, and establish L. occidentalis as a potentially serious pest in lodgepole pine seed orchards.     

The first record on Leptoglossus occidentalis (Hemiptera: Coreidae) in Korea,a potential pest of the pinaceous tree species

Based on body appearance and COI gene sequence, we report a leaf-footed bug, Leptoglossus occidentalis (Hemiptera: Coreidae), for the first time from Korea. The bug was first encountered in Changwon-si, Gyeongsangnam-do, Korea in 2010, and then again in 2011. We should be alert against the species, because it is an important pest of Pinaceae in Europe. Explanations on its brief morphology and photographs are provided. The worldwide distribution and life history of the bug, and its damage symptom are introduced briefly herein.     

Life table for Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae) and prediction of damage in lodgepole pine seed orchards

Abstract 1 Fecundity, egg hatch and nymphal survivorship of Leptoglossus occidentalis Heidemann were determined and used to develop a damage prediction formula for seed bugs in lodgepole pine seed orchards. 2 A single overwintered L. occidentalis female may lay up to 80 eggs during early and mid‐season cone development. 3 Up to 83.4% of eggs that were attached to branches with tape hatched successfully, compared with 53.5% of eggs attached to needles with wire. Egg parasitism was only observed in egg masses attached with wire. 4 Egg masses attached with wire suffered approximately 30% mortality from parasitism, primarily by Gryon pennsylvanicum (Ashmead). 5 Approximately 15.1 and 2.5% of first instars survived to adulthood in 2000 and 2001, respectively. Survival of caged nymphs that were protected from predators was 14.5 and 17.8%, respectively. 6 A composite survival table indicated that 9.3% of eggs laid by L. occidentalis are expected to survive to adulthood. 7 Based on previously determined damage estimates for each life stage of feeding during three periods of cone development, a hypothetical density of one seed bug per tree early in the season will result in an expected seed loss of approximately 310 seeds.     

Distribution analysis of Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae) in South Korea using climate and host plant ensemble maps

Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae) is a notorious pest that causes irreversible damage to coniferous forests in South Korea. However, an effective control strategy is still under development. In this study, we used CLIMEX to analyze the potential distribution of L. occidentalis in relation to climate and host plants in South Korea for the identification of effective control spots. The climate needs of L. occidentalis under current and future climate conditions were analyzed and projected on a map along with the distribution of coniferous forests. The CLIMEX model projected that the area of L. occidentalis distribution would decrease slightly in 2060 compared with that at present. However, it was projected that occurrence in mountainous regions would be sustained, suggesting continued damage to coniferous forests in South Korea.     

New occurrence of the invasive alien leaf‐footed bug Leptoglossus gonagra (Hemiptera: Coreidae) in South Korea

The invasive alien leaf‐footed bug Leptoglossus gonagra (Fabricius 1775), native to the Americas, is herein recorded from South Korea for the first time. Forty nymphs and adults were collected on 22 September 2016 from arable lands of two separate sites. These individuals were identified based on the analysis of morphological characters and cytochrome oxidase c subunit 1 sequences. A neighbor‐joining tree indicated the existence of two clades: Group A from Brazil, and Group B from Asia and Oceania. Mean genetic distance between clades was 8.11%, range 7.60–8.57%. Within clades, mean genetic distance was 0.42% (0.17–0.83%) in Group A and, 0.45% (0.00–0.83%) in Group B. Colonies of 2nd‐final instar nymphs and adults were found, on top of leaves, flowers, fruits, and between the stems of pumpkins (Cucurbita moschata). Nymphs and adults damaged host plants by sucking their leaves, stems, fruits, and buds. After the first discovery in 2016, we were unable to find the bugs again (in 2017–2019). Therefore, the invasion of this species seems to have occurred only temporarily, during the warm season. We suggest that the species may have failed to overwinter as it is normally distributed in regions warmer than Korea. Nevertheless, possible appearances of this species must be continuously monitored as it is a potential pest of various plants cultivated in South Korea.     

Scent gland apparatus in the Western conifer seed bug Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae)

The coreid Leptoglossus occidentalis is a Nearctic bug responsible for severe seed losses to pine orchards. When disturbed, adults and nymphs emit a defensive secretion deemed an allomone. Here we describe the gross morphology of the scent gland apparatus and the related evaporatory structures in nymphs and adults of L. occidentalis, through light microscopy and scanning electron microscopy. Adults of both sexes possess a metathoracic scent gland complex (MTG) including a central orange‐yellow reservoir and a pair of white lateral glands, connected by ducts to the reservoir. The MTG belongs to the diastomian type, with two ostioles located on the metathorax associated with a microsculptured cuticular accumulation area, i.e. evaporatory area, which can prevent the spread of the secretion on to non‐evaporative cuticle and increase scent fluid evaporation. A high number of male‐specific sternal gland pores were observed. These pores and associated glands are likely the source of an attractant pheromone, which could be extremely useful in monitoring and combating this invasive pest. In nymphs, MTG is replaced by two dorsal abdominal scent glands (DAGs) located between the 4th and the 6th urotergites. DAGs are reddish cuticle‐lined sacs with gland cells forming the gland wall; the scent substances are released through two orifices lying on the mid‐dorsal abdominal line between urotergites IV–V and V–VI. Also in nymphs, peculiar cuticular evaporatory areas surround both orifices.     

喙副黛缘蝽初步研究

副黛缘蝽Paradasynus longirostris Hsiao隶半翅目缘蝽科,是莆田龙眼一种重要害虫。该虫50-60年代经调查已见发生危害．但迄未对其进行研究。本是首欢报道。喙副黛缘蝽成虫、若虫刺吸危害龙眼果穗、幼果及叶片。一年发生2代．以成虫越冬;翌年4月下旬出蛰,产卵于叶片上;每雌产卵约60粒,1-2卵块;卵期6-10天。若虫5龄．若虫期20-40天。     

Responses of native egg parasitoids to the invasive seed bug Leptoglossus occidentalis

1. The Western conifer seed bug Leptoglossus occidentalis, a native insect of North America, was accidentally introduced in Europe in the late 1990s. Since then, it has spread rapidly. Biological control could provide an efficient management option but natural enemies of the pest have been poorly examined in Europe.
2. In this study, we exposed sentinel egg masses and collected naturally laid egg masses in southern France in 2016 and 2017, to identify the egg parasitoids of L. occidentalis and investigate their potentials.
3. Three egg parasitoids were detected: Anastatus bifasciatus, Ooencyrtus pityocampae and Ooencyrtus telenomicida. The overall parasitism was low compared to that observed in the native range with 6.4% of all eggs being parasitized, while 17.1% of egg masses carried at least one parasitized egg. The total number of parasitized egg masses was similar between parasitoid species, but the mean number of parasitized eggs per egg mass was highest for A. bifasciatus (5.57 vs. 1.25 for Ooencyrtus spp.).
4. Sentinel egg masses underestimated the parasitism compared to natural egg masses (respectively, 1.42% and 7.71%).
5. Our results suggested that the three generalist parasitoids detected can respond in a Leptoglossus egg density-dependent manner, but this requires further investigations.

     

Revision of the Neotropical genus Nyttum (Hemiptera: Coreidae)

Abstract The Neotropical genus Nyttum, belonging to the wholly New World tribe Acanthocephalini of the plant-feeding heteropterous family Coreidae, is redescribed. Its distribution is shown to extend from Mexico to Uruguay. Nyttum limbatum Spinola, from Brazil, and N.punctatum (Dallas), widespread in South and Central America, are redescribed; Anasa perfusa Distant is synonymized with N.punctatum. Seven new species are described: Nyttum amaralae, from Uruguay; N.beckeri, from Brazil and 'West Indies' N.nigrum, from Venezuela; N.pallens, from Mexico; N.pulchrum, from Peru; N.rubrum, from Brazil; and N.spinolai from Brazil and Guyana. A key to the species is included. Placoscelis is regarded as the genus most closely related to Nyttum and it is shown that the same two types of paramere and the same two types of aedaegus occur in both genera.  

Resumen

. Se redescribe el género neotropical Nyttum, perteneciente a la tribu Acanthocephalini que se encuentra exclusivamente en el Nuevo Mundo, dentro de la familia Coreidae (Hemiptera-Heteroptera) que se alimenta exclusivamente de vegetales. Se indica que la distribución de este género se extiende desde Mexico al Uruguay. Se redescriben Nyttum limbatum Spinola del Brasil y N.punctatum (Dallas), de amplia distribución en América del Sur y Central. Anasa perfusa Distant es sinonimizada con N.punctatum. Se describen siete neuvas especies: Nyttum amaralae, del Uruguay; N.beckeri, del Brasil e 'Indias Occindentales' N.nigrum, de Venezuela; N.pallens, de Mexico; N.pulchrum, de Perú; N.rubrum, de Brasil; y N.spinolai, de Brasil y Guyana. Se incluye una clava para la separation de las especies. Se considera que Placoscelis es el género mas cercano a Nyttum, y se indica que los mismos dos tipos de parameros y los mismos dos tipos de aedeagus se encuentran en ambos.     

缘蝽科四种昆虫同工酶鉴别（半翅目：缘蝽科）

本研究运用聚丙烯酰胺垂直板电泳技术对缘蝽科4个种的酯酶同工酶进行了检测,酯酶酶谱可分为4个区,酯酶酶谱在属间表现明显差异,在属内具有某些共性,聚类分析显示4个种的分类地位,并说明性别差异大于个体差异但小于种间差异。     

Functional anatomy of the vagina muscles in the adult western conifer seed bug,Leptoglossus occidentalis (Heteroptera: Coreidae), and its implication for the egg laying behaviour in insects

The anatomy of the female reproductive tract and the nerve-evoked contractions of the vagina muscles and their association with the ovipositor in the western conifer seed bug, Leptoglossus occidentalis (Heidemann) are investigated for the first time. The reproductive tract consists of a set of paired telotrophic ovaries, each containing seven ovarioles, located in the anterior lateral regions of the abdomen. Each ovary is attached to a lateral oviduct which spans most of the abdomen to attach to a relatively short common oviduct that joins the vagina near the rear of the animal. The vagina is associated with a pair of bilaterally symmetrical muscles attached at their posterior ends to lateral extensions of sternite VIII, the valvifer of the Type II ovipositor. From this attachment site, the muscles fan out medially and anteriorly to converge along the dorsal midline of the vagina up to the base of the common oviduct. Vagina muscles respond to a single stimulation of their motor nerves by producing a smooth contraction lasting approximately 1 s. With increasing frequencies of stimulation, the muscle contractions summate to create a tetanic response. The muscles are fatigue resistant being able to maintain the same degree of tension for up to 10 min at 10 Hz stimulation. Visual observation shows that other muscles associated with the valves of the ovipositor behave in a similar fashion to that of the vagina muscles from which the tension recordings were obtained. Fatigue-resistant vagina muscles are discussed in relation to copulation, sperm transport and this insect's ability to deposit a series of eggs directly onto the surface of a conifer needle in a manner by which eight or more blunt-ended eggs are packed end-to-end in a single row.     

Morphological and biological characteristics of the Korean population of Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae), an invasive insect pest of conifer cones

In this study, we investigated the morphological and biological characteristics of a serious insect pest of conifer cones, the western conifer seed bug (WCSB), Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae), in Korea. WCSB adults shared external morphology with overseas populations, including zig-zag fascia on the coria, dilation of the hind tibiae, and color pattern on the abdominal dorsum. The average body length and width of WCSB adults were 17.20 and 4.85 mm, respectively, in males and 19.05 and 5.55 mm in females. The average body weights were 111.27 and 169.89 mg in adult males and females, respectively. On average, eggs were 2.12 mm long and 1.17 mm wide. The average body lengths of the first- to fifth-instar nymphs were 3.07, 4.76, 8.77, 13.56, and 15.53 mm, respectively. For biological characteristics, developmental durations of the egg and five nymphal instar stages were 8.6, 3.7, 7.6, 5.8, 5.8, and 9.1 days, respectively, in laboratory conditions. Nymphal mortality was 47.5% in total. The average life expectancy of WCSB adults was 58.2 and 63.9 days in males and females, respectively. Oviposition of the adult females started 23.4 days after emergence, and the total number of eggs laid by a single adult female was 68.6 on average. The results of this study offer basic morphological and biological information that can be used to design an effective control strategy to manage the WCSB population in Korea.     

Phylogenomic analysis suggests Coreidae and Alydidae (Hemiptera: Heteroptera) are not monophyletic

Next‐generation sequencing technologies (NGS) allow systematists to amass a wealth of genomic data from non‐model species for phylogenetic resolution at various temporal scales. However, phylogenetic inference for many lineages dominated by non‐model species has not yet benefited from NGS, which can complement Sanger sequencing studies. One such lineage, whose phylogenetic relationships remain uncertain, is the diverse, agriculturally important and charismatic Coreoidea (Hemiptera: Heteroptera). Given the lack of consensus on higher‐level relationships and the importance of a robust phylogeny for evolutionary hypothesis testing, we use a large data set comprised of hundreds of ultraconserved element (UCE) loci to infer the phylogeny of Coreoidea (excluding Stenocephalidae and Hyocephalidae), with emphasis on the families Coreidae and Alydidae. We generated three data sets by including alignments that contained loci sampled for at least 50%, 60%, or 70% of the total taxa, and inferred phylogeny using maximum likelihood and summary coalescent methods. Twenty‐six external morphological features used in relatively comprehensive phylogenetic analyses of coreoids were also re‐evaluated within our molecular phylogenetic framework. We recovered 439–970 loci per species (16%–36% of loci targeted) and combined this with previously generated UCE data for 12 taxa. All data sets, regardless of analytical approach, yielded topologically similar and strongly supported trees, with the exception of outgroup relationships and the position of Hydarinae. We recovered a monophyletic Coreoidea, with Rhopalidae highly supported as the sister group to Alydidae + Coreidae. Neither Alydidae nor Coreidae were monophyletic; the coreid subfamilies Hydarinae and Pseudophloeinae were recovered as more closely related to Alydidae than to other coreid subfamilies. Coreinae were paraphyletic with respect to Meropachyinae. Most morphological traits were homoplastic with several clades defined by few, if any, synapomorphies. Our results demonstrate the utility of phylogenomic approaches in generating robust hypotheses for taxa with long‐standing phylogenetic problems and highlight that novel insights may come from such approaches.     

The population ecology of Amorbus Dallas (Hemiptera: Coreidae) species in Australia

Species of Amorbus Dallas (Hemiptera: Coreidae) appear to exhibit a tight link between oviposition preference and nymphal performance: females apparently select vigorous plant modules to facilitate the development of their offspring. Such behaviour suggests that these insects should exhibit population dynamics consistent with the latent classification proposed by Price et al. (1990). While this empirical hypothesis is intuitively appealing it does not appear to incorporate current population dynamics theory. Berryman's ecodynamics (1992) offers the potential to describe a species' population dynamics which is grounded in a quantitative theoretical framework. The population ecology of Amorbus species from Australia is considered in relation to these hypotheses.     

Comparison of management strategies for squash bugs (Hemiptera: Coreidae) in watermelon

Two watermelon pest management practices, a squash trap crop and a standard recommendation using soil-applied carbofuran, were compared using large-scale field plots to assess trap crop suitability as a replacement for the standard in 2000, 2001, and 2002. In both systems, foliar insecticide applications were used to control squash bugs when populations exceeded threshold levels. During 2001 and 2002, a treatment of untreated watermelon was used. Early season adult insects, from seedling to fruit set, are most critical for watermelon. Significantly fewer early adult bugs were found on watermelon in the trap crop than in the standard recommended practice in 1 of 3 yr. In both years, significantly fewer adult squash bugs were found in watermelon in the trap crop than in untreated fields. The standard recommended practice significantly reduced adult squash bugs in watermelon compared with the untreated in 1 of 2 yr. There was no significant correlation of watermelon yield and squash bug density, indicating that squash bug densities were too low to impact yield. Although squash bugs were reduced significantly by the trap crop, marketable watermelon yields were lower in the squash trap crop than in untreated watermelon, suggesting that pest management treatments may interfere with crop productivity factors other than squash bug colonization. Results suggest that mid-season production squash bug should be managed by monitoring populations and using insecticides as needed rather than using at-plant treatment. Further research is needed to compare treatments during early-season production.     

First records of the western conifer seed bug Leptoglossus occidentalis Heid. (Heteroptera, Coreidae) from Russia and Ukraine, regularities in its distribution and possibilities of its range expansion in the palaearctic region

The paper presents the first records of the western conifer seed bug Leptoglossus occidentalis Heidemann, 1910 for the fauna of Russia (Rostov Province) and Ukraine (Crimea and Zaporizhia Province). The dependence of the spread of this species on positive and negative temperatures and the availability of food plants is considered. The forecast of further expansion of its range in the Palaearctic Region is given.