Notes on the prey,nesting behaviour and natural enemies of three Bembix sand wasps (Hymenoptera: Crabronidae) in the Iberian Peninsula

Bembix is a cosmopolitan genus with more than 300 species distributed all over the world. The females of these wasps dig burrows in the ground and hunt for flies to feed their larvae. Current knowledge of the nesting behaviour of most Palaearctic species (around 50) is limited. In this study, we provide data on the nest structure, nesting activity, natural enemies and prey captured by three Bembix sand wasps present on the Iberian Peninsula: B. zonata, B. flavescens bolivari and B. merceti. We also analyze the prey spectrum of the three species, comparing our data with those available from previous studies. B. zonata exhibits the greatest selectivity index (and the narrowest niche width), capturing mainly Syrphidae to provision its nests. B. merceti shows the greatest niche width in comparison with the other two species. The inter-annual differences in the diet of B. merceti are important and are probably due to the ability of this wasp to change from one prey resource to another, depending on its availability.     

Behavior during artifically induced swarm emigration in an old world polistine wasp,Ropalidia romandi (Hymenoptera: Vespidae)

Absconding swarms of the Australian swarm-founding wasp,Ropalidia romandi, were artificially induced. Workers may initially search for new nesting sites and once a new nest site is decided, they recruit their nest mates including multiple queens there. During the swarm emigration, workers often land on prominent objects, such as tree leaves, and drag the metasoma; those that do not drag usually antennate the object surfaces. These findings suggest thatR. romandi workers use a scent trail during swarm emigration. Artificial transfer of colonies resulted in clustering of wasps from different colonies, indicating thatR. romandi may not act exclusively against non-nestmates. In the formation of tentative aggregation of wasps prior to commencing emigration, and also in recognizing a new nesting site by following swarm members that have been guided by a scent trail, a visual cue (aerial swarm formed around an aggregation of wasps) seems to play an important role.     

Pulp partitioning and worker specialization in polistine wasps (Hymenoptera,Vespidae, Polistinae)

The character of pulp partitioning was studied in three Palaearctic polistine wasps (Polistes dominula and P. gallicus in the Black Sea Biosphere Reserve, and Polistes nimpha in the Lugansk Nature Reserve, Ukraine) and in five Neotropical species (P. lanio, P. versicolor, Mischocyttarus injucundus, M. alfkeni, Metapolybia cingulata, and Protopolybia exigua in Trinidad). In independent-founding Polistes and Mischocyttarus species, all cases of pulp partitioning were connected with aggression on the part of the foundresses. An experimental increase of nesting density of P. dominula led to changes in the relative abundance of workers in the polyethic task groups. Some workers practically stopped hunting and switched over to pulp delivery; some of these workers possessed well-developed ovaries and were able to lay eggs. The experimentally removed part of the nest envelope in the swarm-founding wasp Metapolybia cingulata was restored only in colonies that included young workers. The specific features of nest building activity in independent-founding and swarm-founding polistine wasps are discussed.     

Nesting Habit and Alarm Pheromones in Polistes gallicus (Hymenoptera, Vespidae)

Alarm pheromones are used in many social insects to elicit an alarm response of nestmates towards disturbances. This chemical alarm channel is especially used by species nesting in closed environments. Polistes gallicus paper wasps typically found their nests on open substrates where visual and vibrational stimuli could be more important than the chemical one to alarm the colony. We investigated, through field bioassays, if, also in this species, workers venom is still the source of alarm pheromones. Our results show that venom volatiles are able to increase the colony defence reaction stimulating more individuals to attack and sting the object of disturbance. Therefore our results suggest the hypothesis that chemical alarm is independent of nesting habit (concealed vs exposed) within the Polistes genus.     

Behavioural notes on the Neotropical parasocial spider wasp Ageniella (Lissagenia) flavipennis (Banks) (Hymenoptera: Pompilidae), with host association

1. Ageniella is a species‐rich group of spider wasps restricted to the New World. Knowledge regarding the behaviour of this genus is based mainly on the Nearctic species, which have been reported to nest solitarily in the soil. This study reports for the first time the nesting behaviour, with host association of Ageniella (Lissagenia) flavipennis Banks. 2. Behavioural aspects on the nesting of A. flavipennis were observed from a study of six nests found in an Atlantic Forest conservation area in São Paulo State, Brazil. Host specimens were collected from a nest, as well as while being carried by an A. flavipennis individual. 3. The present study reports the A. flavipennis females cohabiting or nesting solitarily in mud nests, indicating that this spider wasp shows some lower level of parasociality. In addition, the spider Enoploctenus cyclothorax (Bertkau) was reported for the first time as host. As has been observed for other Ageniellini, females of A. flavipennis amputate the host's legs and transport the spider to the nest, flying or walking forward. 4. Communal behaviour has been reported for species of different genera of Pompilidae, such as Macromeris Lepeletier, Paragenia Bingham and Auplopus Spinola. Recent phylogenetic analyses suggest that the Lissagenia species are more closely related to the other Ageniellini genera than to the other Ageniella species. The present information on nesting and prey could contribute towards a more conclusive phylogenetic position of Lissagenia.     

Evolution of nesting behaviour and kleptoparasitism in a selected group of osmiine bees (Hymenoptera: Megachilidae)

The construction of nests to rear offspring is restricted to vertebrates and few insect taxa, such as termites, wasps, and bees. Among bees, species of the family Megachilidae are characterized by a particularly high diversity in nest construction behaviour. Many megachilid bees nest in excavated burrows in the ground, others place their brood cells in a variety of above‐ground cavities or attach them to the surface of a substrate, and yet others have adopted a kleptoparasitic habit. Evolutionary transitions between the different nesting sites and between conventional nesting and kleptoparasitism in bees are poorly understood. In the present study, we traced the evolution of nesting site selection and kleptoparasitism in the Annosmia–Hoplitis group (Osmiini), which displays an exceptionally high diversity in nesting behaviour. We found that the evolution of nesting behaviour proceeded unidirectionally from nesting in excavated burrows in the ground to nesting in rock depressions and cavities, followed by the colonization of snail shells and insect borings in dead wood or hollow stems. Kleptoparasitism evolved once and the kleptoparasitic species have derived from the same lineage as their hosts. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 349–360.     

Nest site competition between cavity nesting passerines and golden paper wasps Polistes fuscatus

Nest boxes provide sheltered nesting sites for both passerines and paper wasps. Although neither wasps nor birds appear to evict the other once one is fully established, it is unclear which is the dominant competitor at the onset of the breeding season. Using wire mesh, we excluded birds but not golden paper wasps Polistesfuscatus from alternating boxes along a transect through edge habitat in North Carolina from 2006 – 2008. If wasps dominate Carolina chickadees Poecile carolinensis and eastern bluebirds Sialia sialis during the early spring (all have similar nest initiation dates), we would expect wasps to settle in both box types at equal frequencies. However, if birds dominate wasps, we would expect wasp nests to be concentrated in “bird‐proof” boxes. We found wasps in bird‐proof boxes significantly more often than in bird‐accessible boxes, indicating that secondary‐cavity nesting birds are able to exclude wasps from available nest sites. Additionally, we found that during the period of nest initiation, birds usurp wasps more often than vice versa.     

Prey selection by a spider wasp,Batozonellus lacerticida (Hymenoptera: Pompilidae): Effects of seasonal variation in prey species,size and density

Prey selection by a spider wasp,Batozonellus lacerticida, was investigated at the riverside of the Toyohira River in Misumai, Sapporo, Hokkaido over 7 years from 1981 to 1987. Seventeen species of araneid spiders were found in the study area and six of them were hunted by the wasps. Most (97.3%) of the 223 prey records obtained were of three species,Araneus macacus, A. marmoreus andA. pinguis. Nesting activity of the wasps continued until mid-August in 1981, 1982 and 1986, but stopped in July in 1984, 1985 and 1987. The length of nesting period was related to whether the wasps successfully switched prey species fromA. macacus toA. marmoreus andA. pinguis andA. pinguis or not Switching seemed to occur at the phase when density ofA. macacus remained high, depending on the density of largerA. marmorus andA. pinguis. This switching had large effects on species composition and size distribution of actual prey. Chesson's index α calculated at every 10 day period revealed that the wasps preferredA. macacus to the two other species and large prey to medium and small ones. The effect of prey density on preference depended on whetherA. macacus was present or absent. In the presence of it there were only slight differences in preference among various conditions of prey density. However, whenA. macacus was absent, densities of bothA. marmoreus andA. pinguis had some complementary effects on preference. The importance of prey size selection by pompilid wasps, and implications of density effects on preference are discussed.     

Developmental time,longevity, reproductive capacity and sex ratio of the mealybug parasitoid Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera: Encyrtidae)

Key biological parameters of Anagyrus sp. nr. pseudococci reared on the citrus mealybug Planococcus citri (Risso) were determined. The mean longevity of female wasps was 21.1 days, the median lethal time (LT₅₀) was 22 days, the mean number of offspring per female was 30.2 wasps and the sex ratio of the progeny was slightly male biased (1.1 males per female). Wasps fed with water showed a significantly lower longevity compared to that recorded for wasps fed with syrup for bee nutrition mixed with water. The longevity of female wasps was significantly higher compared to males when the syrup-water solution was provided as food. Longevity was significantly higher for female wasps fed with syrup and water in comparison with those exposed to host mealybugs. The mean development time of female and male wasps was 14.7±0.1 and 14.5±0.1 days, respectively.     

Do digger wasps time their provisioning activity to avoid cuckoo wasps (Hymenoptera: Crabronidae and Chrysididae)?

Strategies adopted by parasitoids and kleptoparasites co-evolve with the defensive adaptations of their hosts, and vice-versa. Hedychrum rutilans and Hedychrum nobile are brood parasites of, respectively, Philanthus triangulum and Cerceris arenaria, two digger wasps that share most aspects of their nesting biology (solitary females dig aggregated nests in the ground and mass-provision the brood with paralyzed insects). We tested the hypothesis that similarity in the hosts’ nesting habits corresponds to similar defensive strategies against these cuckoo wasps. Peak provisioning activity by P. triangulum occurred in late afternoon (and early morning in 1 year) while peak H. rutilans activity was in early afternoon. In contrast, peak provisioning by C. arenaria and peak H. nobile activity occurred in early afternoon. Thus, P. triangulum (as previously found in other populations) appears to have timed its provisioning to avoid its brood parasite whereas C. arenaria did not, rejecting our hypothesis. The daily activity of both chrysidid wasps was positively correlated to air temperature. Host nest density positively affected only H. rutilans activity, in agreement with previous reports for other populations of this species, whereas the daily pattern of activity of the host was the key correlate with the activity of H. nobile. Mortality due to cuckoo wasps was low for both digger wasps, although it was somewhat higher at sites with many nests of P. triangulum. We suggest that perhaps the degree to which digger wasps time their activity to avoid cuckoo wasps is related to the degree of specialization in host choice by their brood parasite. H. rutilans is more highly dependent on P. triangulum, which may have had a greater effect on the timing of provision by its host.     

Nest-site selection in the great golden digger wasp, Sphex ichneumoneus L. (Sphecidae)

Abstract. 1. Sphex ichneumoneus are solitary, ground-nesting wasps with apparently broad nesting requirements.
2. Nest-site selecting behaviour involves four levels of decisions. First, the wasps are not active unless temperatures are high and they are less active during cloudy weather. Second, at the beginning of a nesting sequence they generally fly to the site from which they emerged, but occasionally they fly to other flat, vegetation-free areas. Third, after landing, they engage in a predigging search of the area, walking around, looking into and briefly digging in the nests of other wasps. In this way they apparently compare and choose to begin digging in the warmest areas available which are not too close to other nests. Fourth, if the soil is too soft and collapses or if it is too hard-packed, the wasp resumes the predigging search. The preference for digging within a range of compactness means that they generally choose sandy loam.
3. Theory would suggest that the wasps may spend an optimal amount of time in the predigging search behaviour and that frequency-dependent selection may be an important factor in the choice of nesting sites.
4. By choosing nesting sites according to these rules, the wasps are ensuring that their offspring develop in the warmest and best-drained soils available.     

Colony Development and Serial Polygyny in the Primitively Eusocial Wasp Belonogaster juncea juncea (Vespidae: Polistinae)

In Cameroon, the social ropalidiine wasp Belonogaster juncea juncea mostly selects human constructions for nesting, but nests installed in the shelter of large boulders were also noted. Nests were founded throughout the year by one female (25.5%) or by associations of two to eight females. Among the 49 colonies studied from foundation, 24 produced at least one adult, and only 5 produced males among the eight that succeeded in producing sexual individuals. After the first dominant female disappeared (died or abandoned the nest, from 77 to 196 days after foundation) the remaining females fought among themselves, leading to a drastic decrease in the total number of brood. They completely abandoned 22 of 24 surviving nests, over a mean colony cycle lasting about 7 months. Hence the colony and nesting cycles were congruent, illustrating that determinate nesting corresponds to the majority of the cases in this species. In the other two nests, some females remained on the nest and began a new colonial cycle illustrating semi-independence between colony and nesting cycles. Up to four successive cycles were completed in this way in the same nest, thus illustrating the phenomenon of serial polygyny (intermediate between determinate and indeterminate nesting cycles). Serial polygyny in independent founding polistine wasps, previously documented only once, has implications for studies attempting to identify factors involved in regulating the colony and nesting cycles in tropical social wasps.     

Biological notes on a fungus-growing ant, Trachymyrmex cf. zeteki (Hymenoptera, Formicidae, Attini) attacked by a diverse community of parasitoid wasps (Hymenoptera, Diapriidae)

A number of wasps in the family Diapriidae, subfamily Diapriinae (Proctotrupoidea), are parasitoids that specialize on ant larvae. These wasps are abundant and diverse in the Neotropics, but little is known about their biology. We studied parasitism rates by an array of diapriine wasps that attack the larvae of fungus-growing ants, Trachymyrmex cf. zeteki, in a single population (near Gamboa, Panamá). Relatively little is known about the biology and natural history of these ants, so we also present data on colony size and nest architecture. We excavated 136 colonies in central Panamá from June to September 2006, and 20 nests from July 2009. We reared six wasp morphotypes; two of them in the genus Mimopriella Masner and Garcia, one Oxypria Kieffer, two Szelenyiopria Fabritius and one Acanthopria Ashmead. The mean intensity of larval parasitism per ant colony was 33.9% (2006), and its prevalence across all ant populations was 27.2% (2006 and 2009). Parasitism rates were not positively correlated with host colony size. A single case of super-parasitism was documented in which two Oxypria males were reared from the same host larva.     

西藏波密柑马蜂的筑巢习性（英文）

柑马蜂Polistes mandarinus在不同的环境中会利用植物材料制作各种不同的蜂巢。我们在2009年5-8月间观察研究了西藏自治区波密县的柑马蜂筑巢习性, 并收集62个蜂巢。柑马蜂利用植物纤维和树脂将它们的巢穴筑于当地居民的木屋檐下。收集的42个在用蜂巢中, 8月末平均拥有79.64±65.28个蜂房, 13.1±10.68个蜂蛹。20个弃用蜂巢中, 平均拥有12.95±3.99个蜂房。8月末, 成功的柑马蜂群拥有约80个个体。研究地的柑马蜂采取3种方式来筑巢, 但以第1种方式最为普遍。第1种筑巢方式筑造的蜂巢以两个蜂房连接巢柄, 结构最为稳定, 所以也应用最广, 后两种筑巢方式筑造的蜂巢均以一个蜂房连接巢柄, 是由于越冬雌蜂在找不到合适的筑巢材料时才采取的建巢方式。     

Monitoring of tube‐nesting bees and wasps with bamboo tube nest traps of different types in two types of forests in temperate Japan

Tube‐nesting bees and wasps were collected with bamboo tube nest traps in a cherry blossom forest and an old secondary forest in Tama Forest Science Garden in the western suburbs of Tokyo. Curtain type traps, in which bamboo tubes were vertically arranged, attracted more bees and wasps to nests than bundle type traps, in which bamboo tubes were bundled haphazardly. The attractiveness of small tubes was similar between small tube traps, which were only composed of small tubes (about 4 mm in diameter), and size mixture traps, which set small tubes together with larger tubes, indicating that the presence of the larger species did not affect the nesting of small species. In the cherry forest, which was characterized by the sparse distribution of deciduous cherry trees with a thin canopy layer and rich growth of weeds, Megachilidae, Colletidae, Sphecidae and Eumenidae were common. In the shady secondary forest, where evergreen trees (Abies firma Sieb. et Zucc., Quercus glauca Thunb.) and shrubs (Aucuba japonica Thunb., Eurya japonica Thunb., Camellia japonica L.) were prevalent, Pompilidae was abundant. Among five species of Crabronidae recorded in this study, four were collected mostly in the cherry forest, whereas the most abundant, Trypoxylon malaisei Gussakovskij, was also common in the secondary forest. Likewise, Auplopus carbonarius (Scopoli) (Pompilidae) was common in the secondary forest and not rare in the cherry forest. These results suggest that the tube‐nesting bees and wasps can be a good bioindicator for monitoring environment.     

Observations on the nesting behavior ofMoniæcera asperata (Fox) (Hymenoptera,sphecidæ, crabroninæ) with comments on communal nesting in solitary wasps

Summary Moniæcera asperata (Fox), a species occuring in southwestern United States and provisioning its nests in the soil with small Diptera and Hemiptera, has been found to exhibit communal nesting, two or three females sharing the same burrow. Similar behavior has been reported for four other Sphecidæ, all five belonging to the tribe Crabronini. Thus both types of social evolution believed to occur in bees, by way of communal nesting and by way matrifilial associations, are exemplified, in their initial stages, in the solitary digger wasps.

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Zusammenfassung Moniæcera asperata (Fox), eine Art von den südwestlichen Vereinigten Staaten, versorgt ihre Erdnesten mit kleinen Diptera und Hemiptera, und hat gemeinschaftliche Nester mit 2–3 Weibchen die denselben Bau teilen. Ähnliches Verhalten ist von vier andere

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Résumé Moniæcera asperata (Fox) est une espèce existant dans le Sud-Ouest des Etats-Unis et qui approvisionne ses nids creusés dans le sol de petits Diptères et Hémiptères. On a découvert chez cette espèce une tendance sociale qui se manifeste par le fait que plusieurs femelles (deux ou trois) partagent le même terrier. Ce comportement a déjà été observé chez quatre autres Sphecidæ, appartenant tous, comme celui-ci, à la tribu des Crabronini. Ainsi, les deux types d'évolution sociale qu'on croit s'être réalisés chez les abeilles, soit par nidification en commun, soit par association des filles avec leur mère, se retrouvent dans leurs phases initiales chez les Hyménoptères fouisseurs.

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Research supported by theNational Science Foundation as part of a comparative study of the behavior of solitary wasps, grant no. G 17497.     

Observations onApoica pallens,a nocturnal neotropical social wasp (Hymenoptera: Vespidae,Polistinae, Epiponini)

Summary Colonies and nests ofApoica pallens in the llanos region of Venezuela range from small foundress nests to large mature colonies. Nests are sited on small diameter, near-horizontal branches in a variety of shrub and tree species. During the day, adult wasps cluster on the face of the nest in an array that seems to be determined by orientation to gravity; defense of the colony against parasitoids and ants by the resting wasps may be more a passive than an active behavior. Wasps fan their wings to cool the colony during the day, but no foraging for water accompanies the fanning behavior. Nightly foraging activity begins with the explosive departure from the nest of hundreds of wasps, most of which rapidly return. Moderate foraging levels early at night give way to very low foraging levels in pre-dawn hours. The period of moderate foraging may be extended for longer hours during increased moonlight. Foraging wasps collect arthropod provisions for larvae. Larvae produce a trophallactic saliva; adults engage in inter-adult trophallaxis; brood are cannibalized. During cluster formation prior to swarm emigration, adult wasps do not appear to scent-mark substrates such as leaves. Instead,A. pallens exhibits a calling behavior, unique among polistine wasps studied to date, in which the gaster is held rigidly away from the thorax and metasomal sternal glands are exposed. Swarms can emigrate during the day.A. pallens may incorporate absconding and colony relocation as features of its colony cycle in the highly seasonal llanos.     

Increase trend of social hymenoptera (wasps and honeybees) in urban areas,inferred from moving‐out case by 119 rescue services in Seoul of South Korea

The number of social hymenoptera have decreased due to accelerated urbanization in Seoul, Korea, during the 1970s–1990s. However, recent changes in environmental conditions have brought distinct re‐growth of their colonies. In this study, we analyzed the re‐colonization trend of social hymenoptera in Seoul with moving‐out reports submitted to 119 rescue services from 2000 to 2009. A total of 14 253 social hymenoptera findings and treatment cases were reported during that time, and the reported numbers increased continuously every year. Among them, 87% of the cases were related to social wasps and 13% were for honeybees. Social wasps peaked from July to September, when the colonies matured, whereas honeybees peaked from April to July when colonies divided. As observed by geographical information system (GIS), moving‐out cases formed hotspots in low‐story buildings around green areas such as nearby forests or city green parks in Eunpyeong‐gu and Gwanak‐gu in Seoul. Among the social wasps, the most frequent species that caused a nuisance and direct stinging was Polistes rothneyi koreanus, the majority of which nested under eaves (63%). Among the honeybees, Apis mellifera was a nuisance while attaching to buildings and walls in the city (60%) during hive splitting. We present herein the situation of the return of social hymenoptera to Seoul and discuss the possible reasons for the recent increase in social wasps in urban areas, including enlarging the green space in urban areas and stable nesting places, which benefited colony development due to the relatively higher urban temperature and the few natural predators and parasitoids.     

Comparison of Buprestidae collected by Cerceris fumipennis (Hymenoptera: Crabronidae) with those collected by purple prism traps

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Some aspects of interrelations between ants (Hymenoptera,Formicidae) and polistine wasps (Hymenoptera,Vespidae)

Embodied, situated and enactive aspects of relationships of polistine wasps with ants are considered within the framework of the theory of autopoiesis. The idea of the embodied interaction implies specific nestbuilding and protective behavior in polistine wasps. The paper examines the adaptive role of applying ant repellent on the petiole and nest and the latitude gradient of such behavior in re-social wasp species. The situated interaction is considered in the environmental context: the mortality of Polistes gallicus (L.) colonies as a result of predatory attacks of ants Myrmica bergi Ruzsky is analyzed in the Lower Dnieper basin (Ukraine). The enactive interaction includes both spontaneous autonomous activity of its participants as a result of self-organization and specific features of the spatial structure of the prey’s population forming under the impact of the predator. The applicability of some “predator-prey” models is discussed.