No abstract

Sociality may affect symbiosis and vice versa. Many plant-sucking stinkbugs harbor mutualistic bacterial symbionts in the midgut. In the superfamily Pentatomoidea, adult females excrete symbiont-containing materials from the anus, which their offspring ingest orally and establish vertical symbiont transmission. In many stinkbug families whose members are mostly non-social, females excrete symbiont-containing materials onto/beside eggs upon oviposition. However, exceptional cases have been reported from two subsocial species representing the closely related families Cydnidae and Parastrachiidae, wherein females remain nearby eggs for maternal care after oviposition, and provide their offspring with symbiont-containing secretions at later stages, either just before or after hatching. These observations suggested that sociality of the host stinkbugs may be correlated with their symbiont transmission strategies. However, we found that cydnid stinkbugs of the genus Adomerus, which are associated with gammaproteobacterial gut symbionts and exhibit elaborate maternal care over their offspring, smear symbiont-containing secretions onto eggs upon oviposition as many non-social stinkbugs do. Surface sterilization of the eggs resulted in aposymbiotic insects of slower growth, smaller size and abnormal body coloration, indicating vertical symbiont transmission via egg surface contamination and presumable beneficial nature of the symbiosis. The Adomerus symbionts exhibited AT-biased nucleotide compositions, accelerated molecular evolutionary rates and reduced genome size, while these degenerative genomic traits were less severe than those in the symbiont of a subsocial parastrachiid. These results suggest that not only sociality but also other ecological and evolutionary aspects of the host stinkbugs, including the host-symbiont co-evolutionary history, may have substantially affected their symbiont transmission strategies. (250 words).

In recent years, three terrestrial bugs, Adomerus triguttulus and Sehirus cinctus (Cydnidae) and the closely related Parastrachia japonensis (Parastrachiidae), have been the focus of several fascinating studies because of the remarkable, extensive parental care they were found to display. This care includes egg and nymph guarding, production of trophic eggs, unfertilized, low cost eggs that are used as food by newly hatched nymphs, and progressive provisioning of the host seed. In this study, we have investigated yet a third related Asian cydnid, Canthophorus niveimarginatus, with regard to the possible occurrence of some or all of these complex traits in order to assess how widespread these maternal investment patterns are in this group of insects and to better understand the implications of their manifestations from an evolutionary context. Manipulative experiments were carried out in the lab to determine whether females provision nests. Observational and egg removal studies were carried out to determine whether trophic eggs are produced, and, if they are, their possible impact on nymphal success. The findings revealed that C. niveimarginatus does, in fact, progressively provision young, and this species also displays all of the other behaviors associated with extended parental care in subsocial insects. Moreover, unlike the other two related species, which place trophic eggs on the surface of the original egg mass, C. niveimarginatus produces both pre- and post-hatch trophic eggs. Nymphs deprived of access to post-hatch trophic eggs had significantly lower body weight and survival rate than those that fed on them. To our knowledge, this is the first time the production of both pre- and post-hatch trophic eggs has been demonstrated in insects outside the Hymenoptera. In this paper, we qualitatively and quantitatively demonstrate the provisioning behavior and patterns of trophic egg production in C. niveimarginatus. When and how trophic eggs are produced and delivered to young should have important correlations with the ecological and life history constraints under which a species has evolved. Thus, we also discuss the possible ecological and life history factors that favor the evolution of post-hatch trophic eggs.

Subsociality involving provisioning for offspring has been reported for one shield bug and three burrower bug species (Hemiptera: Pentatomoidea: Parastrachiidae and Cydnidae: Sehirinae). We present the first report of subsocial behaviour in the burrower bug Adomerus variegatus (Signoret), focusing on manifestations of maternal care, specifically the production of trophic eggs, guarding of offspring, and progressive provisioning. In our study, each female produced an egg mass that included some trophic eggs. Prior to nymphal hatch, females remained in their nests and showed egg mass guarding behaviour. Mothers started provisioning behaviour a few hours after nymphal hatch. Each mother dragged a seed (larger than her body) with her proboscis to the nest as food for the nymphs. These findings suggest that A. variegatus shows maternal-care behaviours similar to those reported in some other pentatomoid species and that trophic egg production and nest provisioning have evolved as interrelated processes.

In contrast to an open environment where a specific celestial cue is predominantly used, visual contrast of canopies against the sky through the gap, known as canopy cues, is known to play a major role for visually guided insect navigators in woodland habitats. In this paper, we investigated whether a subsocial shield bug, Parastrachia japonensis, could gauge direction using canopy cues on a moonless night. The results show that they could perform the round trip foraging behaviour even in an experimental arena with only an artificial round gap opened in the ceiling of the arena and adjust their homing direction for a new azimuth when the gap was rotated. Thus, P. japonensis can use slightly brighter canopy cues as a compass reference but not complex landmarks during nocturnal homing behaviour.