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The highly polyphagous and invasive brown marmorated stink bug, Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), has become a significant insect pest in North America since its detection in 1996. It was first documented in northern Utah in 2012 and reports of urban nuisance problems and plant damage have since increased. Biological control is the preferred solution to managing H. halys in North America and other invaded regions due to its alignment with integrated pest management and sustainable practices. Native and non-native biological control agents, namely parasitoid wasps, have been assessed for efficacy. Trissolcus japonicus (Ashmead) (Hymenoptera: Scelionidae) is an effective egg parasitoid of H. halys in its native range of southeast Asia and has recently been documented parasitising H. halys eggs in North America and Europe. Field surveys for native and exotic egg parasitoids using wild (in situ) and lab-reared H. halys egg masses were conducted in suburban and agricultural sites in northern Utah from June to September 2017–2019. Seven native wasp species in the families Eupelmidae and Scelionidae were discovered guarding H. halys eggs and adult wasps from five of these species completed emergence. Native species had low mean rates of adult emergence from wild (0.5–3.7%) and lab-reared (0–0.4%) egg masses. In 2019, an adventive population of T. japonicus was discovered for the first time in Utah, emerging from 21 of the 106 wild H. halys egg masses found that year, and none from lab-reared eggs. All T. japonicus emerged from egg masses collected on Catalpa speciosa (Warder). Our results support other studies that have observed biological control of H. halys from T. japonicus and improved parasitoid wasp detection with wild as compared to lab-reared H. halys egg masses.

Halyomorpha halys is a severe invasive Asian pest worldwide and classical biological control is foreseen as the most promising control method. Egg parasitoids appear to be the most important natural enemies of this pest, especially the Asian hymenopteran Trissolcus japonicus. In the invaded areas, only a few egg parasitoid species have been able to adopt H. halys as a host. Anastatus bifasciatus is the most common native egg parasitoid of H. halys in Europe, but reaches only low levels of parasitization, while several other native species are only occasionally found. Recently, adventive populations have been found both in the USA and in Europe of T. japonicus, and in Italy of a second Asian species, Trissolcus mitsukurii. Species identification based on morphological traits by specialists or by molecular analysis is a crucial step in the management of biological control programs. The ability to identify the genus or species within a narrow guild of egg parasitoids based on adult emergence holes and meconium features can be a simple and useful method to support management efforts. We present here detailed descriptions of the meconium of the most frequent parasitoid species attacking H. halys in Europe and the characteristics of their emergence holes of the adult wasps.

As the brown marmorated stink bug (Halyomorpha halys) has spread across the Northern Hemisphere, research on its egg parasitoids has increased accordingly. These studies have included species-level taxonomy, experimental assessments of host ranges in quarantine, and surveys to assess parasitism in the field. We here present a molecular phylogeny of Trissolcus that includes all species that have been reared from live H. halys eggs. Species-group concepts are discussed and revised in the light of the phylogenetic analyses. The analyses indicate that the ability to successfully parasitize H. halys eggs is not phylogenetically constrained, but the most effective parasitoids are all found in the flavipes species group.

The invasive and highly polyphagous brown marmorated stink bug, Halyomorpha halys (Stål), is a severe agricultural and urban nuisance pest in North America. Since its initial invasion into Utah in 2012, H. halys has become well established in urban and suburban locations along the western foothills of the Wasatch Front in northern Utah. Bordering the Great Basin Desert, this area is unique from other North American locations with H. halys due to its high elevation (> 1200 m), aridity (30-year mean RH = 53.1%; dew point = -1.9 °C) and extreme temperatures (the 30-year mean minimum and maximum in January and July in Salt Lake City range from -3.1 to 3.6 °C and 20.3 to 32.4 °C, respectively). To document which plant species harbour H. halys, surveys were conducted in 17 urban/suburban sites in four counties during 2017 and 2018. Halyomorpha halys was more abundant in Salt Lake and Utah counties than in the more northern counties of Davis and Weber and was found on 53 plant species, nine of which hosted two or more developmental stages in both years. The majority of hosts were in the families Fabaceae, Rosaceae and Sapindaceae. Northern catalpa, Catalpa speciosa (Warder), was the most consistent host, supporting a majority of H. halys detections in all life stages; thus we identify it as a sentinel host. Twenty-nine species were novel hosts for H. halys in North America; of these, Acer ginnala Maxim, Populus tremuloides Michx., Prunus armeniaca × domestica ‘Flavor King’ and Prunus virginiana ‘Schubert’ were detected with two or more life stages of H. halys in both years. Peak populations of H. halys occurred from mid-June to mid-September. We describe H. halys plant utilisation by life stage and seasonal period to aid future detection and management of this invasive insect in the greater Intermountain West region.

Uradenia, often referred to as paragenital glands, are usually voluminous paired exocrine glands located ventrally in the abdomen mostly on the intersegmental membrane between abdominal segments (= urites) VII-VIII or VIII-IX, depending on sex or the taxon. They have been previously recorded from eight pentatomomorphan families belonging to Coreoidea, Lygaeoidea and Pyrrhocoroidea (Hemiptera: Heteroptera), found either in males, females or both sexes, and were thought to be absent in Pentatomoidea. We report here the first instance of uradenia in a pentatomoid genus, the African Dismegistus Amyot & Serville, 1843 (Parastrachiidae). Only the male adult possesses uradenia located on the intersegmental membrane of segments VIII-IX. The only other genus of the family, Parastrachia Distant, 1883, as well as other examined genera belonging to pentatomoid families possibly related to Parastrachiidae (Cydnidae, Thyreocoridae), do not possess uradenia. The uradenia of Dismegistus exhibit the same fundamental structure as in other trichophoran families but differ by their dorso-lateral position (instead of ventral), and also by the paired orifices (instead of unpaired and median). The implications of the presence of uradenia within member of a pentatomoid genus are briefly discussed.

A parasitoid wasp, Trissolcus basalis (Wollaston), was recorded parasitizing eggs of the invasive stink bug Halyomorpha halys (Stål) in the United States. This is the first record of this species parasitizing fresh and frozen eggs of H. halys in the United States. First record of Trissolcus basalis parasitizing Halyomorpha halys eggs in the United States.

Cuticular parts of the spermatheca and associated vaginal structures (chiefly the ring sclerites of the parietovaginal glands) have been examined and compared in 190 cydnid species representing 65 genera and all five subfamilies currently recognized in the family (Amnestinae, Cephalocteinae, Cydninae, Garsauriinae, Sehirinae). Four species belonging to genera formerly included within the Cydnidae (Dismegistus, Parastrachia, Thaumastella, Thyreocoris) were also examined. Morphology of the three main parts of the spermatheca (seminal receptacle (distal bulb), intermediate part (pump apparatus), spermathecal duct) is described. Four main types of spermathecae can be recognized from the distal receptacle and the intermediate part: the amaurocorine type (in Sehirinae: Amaurocorini), amnestine type (in Amnestinae), garsauriine type (in Garsauriinae), and "cydnoid" type (in Cephalocteinae + Cydni- nae: Cydnini, Geotomini + Sehirinae: Sehirini). No synapomorphy of these types was found which suggests that the currently con- ceived Cydnidae are not monophyletic. Moreover, out of these four types only the "cydnoid" is typically pentatomoidean due to the presence of an intermediate part usually well delimited by two flanges and having always an unsclerotized flexible zone as well as two internal cuticular structures (septum and fretum) partly obstructing the lumen. The simple tubular amaurocorine type is unusual and aberrant within all Pentatomoidea. The amnestine and garsauriine types display some similarities with taxa outside the Pentato- moidea, especially with some lygaeoid or coreoid spermathecae, mainly in the structure of the intermediate part not delimited proxi- mally (absence of flanges) and devoid of the flexible zone. Within the "cydnoid" type, six spermathecal facies can be characterized principally according to the shape of both the apical reservoir along with the intermediate part, and the differentiations of the sper- mathecal duct. It has been impossible to find any synapomorphy for all species and for the six facies belonging to the "cydnoid" type of spermatheca. We suggest that the Cydnidae as defined presently are probably a polyphyletic group; moreover its main "cydnoid" branch, called by us Cydnidae sensu stricto (Cephalocteinae + Cydninae + Sehirinae: Sehirini) seems to be relatively recent among the Pentatomoidea. Nishadana and Nishocoris are transferred from Garsauriinae back to Cydninae: Cydnini and the tribe Amauro- corini (Sehirinae) is upgraded to a separate subfamily Amaurocorinae stat. nov. Moreover, we regard the Geotomini and the Sehirini both as non-monophyletic and we indicate that by appending them sensu lato (Geotomini "s. l.", Sehirini "s. l.") Contents

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Cydnidae are defined as those Pentatomoidea with setal combs at the apices of the coxae and a strigil on the ventral surface of the metathoracic wing. Eight subfamilies are recognized: Thaumastellinae, Amnestinae, Cydninae, Scaptocorinae, Sehirinae, Garsauriinae, Thyreocorinae (including only Thyreocoris, Carrabas and Strombosoma) and Corimelaeninae. Eumetopia is included in Corimelaeninae. Dismegistus is excluded from Cydnidae.