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.     

Rubbing behavior and morphology of van der Vecht's gland inBelonogaster petiolata (Hymenoptera: Vespidae)

Females of the social wasp, Belonogaster petiolata,rub the secretion of van der Vecht's gland, located on their terminal gastral sternite, onto the nest pedicel. In bioassays, the secretion was repellent to two species of ants, while shortchain acids were effective releasers of rubbing behavior. Rubbing was associated with pedicel enlargement and departure from the nest in preemergence colonies. Its frequency was high where wasps were often exposed to ants and low where ants were rare or absent. Rubbing also decreased significantly from the pre-to the postemergence stage of the colony cycle. In both stages, subordinate foundresses rubbed more often than queens or workers. These observations support the hypothesis that rubbing behavior and the secretion of van der Vecht's gland function in chemical defense of the nest against ant predation. The general morphology of the gland in B. petiolataresembles that of the four other independent-founding polistine wasp genera.     

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.     

Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on Banisteriopsis malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system

In the Brazilian savanna many plant species bear regular associations with patrolling ants that are aggressive towards insect herbivores. However, not only ants but also several species of predatory wasps are attracted to plants due to the extrafloral nectaries (EFNs). Such wasps feed on both herbivores and plant exudates. In this study we describe the foraging behavior of the social Polistinae wasp Brachygastra lecheguana in the extrafloral nectaried shrub Banisteriopsis malifolia, and investigated the influence of patrolling ants Camponotus blandus on the activity of the wasp. Brachygastra lecheguana fed on the endophytic larvae of Anthonomus (Curculionidae) beetles that developed inside flower buds. The wasp lacerated the bud layers to reach the beetle larvae located at the bud core. The wasp visits to Ba. malifolia were statistically related to the abundance of flower buds and beetles. Ant exclusion experiments revealed that the hunting behavior of B. lecheguana on beetles was not related to the absence of C. blandus. However we found that wasps spent more time consuming extrafloral nectar on branches where ants were excluded. This is the first study reporting extrafloral nectar consumption by B. lecheguana, as well as the predation on herbivores in natural areas. In cerrado vegetation, ants benefit the plant by reducing insect herbivores, and our study provides evidence that the B. lecheguana – Ba. malifolia system represents a potential interaction where the wasp may also benefit the host plant. The value of this wasp species as a plant‐guard is discussed.     

Host acceptance behavior of the Japanese aquatic wasp,Agriotypus gracilis (Hymenoptera: Ichneumonidae) toward the caddisfly host,Goera japonica (Trichoptera: Limnephilidae)

The host acceptance behavior of the Japanese aquatic wasp,Agriotypus gracilis Waterston, an ectoparasitoid of the sand case building caddisfly,Goera japonica Banks was investigated in the laboratory. Female wasps were observed to enter the water by walking down a stone protruding from the water surface. Antennae were held backward and not utilized in searching for hosts under the water. Female wasps examined hosts from the outside of their cases by 2 consecutive steps, “turning” and “probing”. Turning behavior, in which female wasps move between the anterior and posterior ends of host cases, may be related to the measurement of case size. Host stages are considered to be discriminated by probing, in which females probe host cases with their sheathing ovipositors. Female wasps most frequently accepted and oviposited on pupal and prepupal hosts.     

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.     

An uneasy alliance:a nesting association between aggressive ants and equally fierce social wasps

Although the Neotropical territorially dominant arboreal ant Azteca chartifex Forel is very aggressive towards any intruder,its populous colonies tolerate the close presence of the fierce polistine wasp Polybia rejecta(F.).In French Guiana,83.33%of the 48 P.rejecta nests recorded were found side by side with those of A.chartifex.This nesting association results in mutual protection from predators(i.e.,the wasps protected from army ants;the ants protected from birds).We conducted field studies,laboratorybased behavioral experiments and chemical analyses to elucidate the mechanisms allowing the persistence of this association.Due to differences in the cuticular profiles of the two species,we eliminated the possibility of chemical mimicry.Also,analyses of the carton nests did not reveal traces of marking on the envelopes.Because ant forager flows were not perturbed by extracts from the wasps’Dufour’s and venom glands,we rejected any hypothetical action of repulsive chemicals.Nevertheless,we noted that the wasps"scraped"the surface of the upper part of their nest envelope using their mandibles,likely removing the ants'scent trails,and an experiment showed that ant foragers were perturbed by the removal of their scent trails.This leads us to use the term"erasure hypothesis."Thus,this nesting association persists thanks to a relative tolerance by the ants towards wasp presence and the behavior of the wasps that allows them to"contain"their associated ants through the elimination of their scent trails,direct attacks,"wing-buzzing"behavior and ejecting the ants.     

Morphology of the fifth sternal glands of Neotropical social wasps (Hymenoptera,Vespidae, Polistinae)

Glands play a central role in the social behavior of insects, and comparative analysis of their structure may provide insights about their evolution and their role in insect social biology. Here, we describe the glands of the fifth metasomal sternite of ten polistine wasps with different social behaviors. The glands of foragers of these species were studied using semithin sections of Araldite‐embedded tissue, and scanning electron microscopy. The structure of the glands observed was mostly similar to what has previously been reported for independent‐founder and swarm‐founder species, respectively. Scale‐shaped modifications in the fifth sternite are reported for the first time for members of the genera Synoeca and Epipona. Two previously unreported glands in the fifth sternite are also described. A more restrictive definition of the name Richards' gland is proposed, applying to the class 3 cells associated with scale modifications of the fifth sternite, based on structural differences between the glands observed, homology requirements, and functional evidence. Previous phylogenetic reconstructions of this character suggest that it appeared early in the Vespidae and disappeared four times, with a single reappearance in the genus Vespula. The restricted definition of Richards' gland suggests that this structure is an exclusive trait of the Epiponini, with two reversals.     

Territoriality in the Malaysian giant ant Camponotus gigas (Hymenoptera/Formicidae)

In a 5-ha area of primary lowland rain forest in Borneo, we observed four polydomous colonies of the night-active giant ant Camponotus gigas. The nonoverlapping, three-dimensional territories in the canopy had a ground size up to 0.8 ha. C. gigas showed a distinct territorial behavior: (1) specific “barrack” nests, especially containing many majors, were situated at the borders and were established during long-term territorial conflicts; (2) trunk trails were regularly patrolled by majors that attacked alien conspecifics and some other ant species violently; and (3) sentinels, often involved in long-enduring conflicts with neighboring ant colonies, defended the borders at bridgeheads. Interspecific conflicts with sympatric Camponotus species always led to violent, “bloody” fights of all castes. Intraspecific conflicts, however, were solved by ritual fights (“front leg boxing”) of majors. C. gigas performed a de-escalation strategy to end short periods of true intraspecific “ant war” that we provoked experimentally. Artificially induced ritualized combats continued over weeks also in the absence of baits, indicating that borders may become established by long-term conflicts of attrition. We discuss the differences between ritual fights in desert and rain-forest ants and apply Lanchester's theory of battles to our findings. Received: October 17, 2000 / Accepted: May 14, 2001     

Oral trophallaxis in adult leaf-cutting ants Acromyrmex subterraneus subterraneus (Hymenoptera, Formicidae)

Adult leaf-cutting ants of the subspecies Acromyrmex subterraneus subterraneus were fed with an Evans Blue dye solution, which allowed the investigation of subsequent exchange of liquids between ants by oral trophallaxis. Trophallactic behavior was filmed and the antennation patterns of donor and recipient ants were described. The ants’ crop capacity was measured following ad libitum feeding on dye solution. Ants previously fed on the dye solution (donors) were placed individually with unfed ants of the same caste (recipients) and the amount of dye solution passed from the donor to the recipient by oral trophallaxis was measured after one hour. The volume received was 0.26 ± 0.15μL (mean ± SD) and the residual volume of dye in the crop of the donors was 0.49 ± 0.23μL. There were 38 trophallactic events recorded for 50 pairs of ants. Trophallaxis was observed from 2.3 min to 21.5 min after initial exposure, with a mean latency of 8.4 ± 5.6 min. The mean duration of a trophallatic event was 2.3 ± 1.3 min. As far as we know, this is the first time that trophallaxis in leaf-cutting ants has been described and quantified. Received 2 December 2005; revised 6 April 2006; accepted 10 April 2006.     

Temporal relationships of rubbing behavior with foraging and petiole enlargement inParapolybia indica (Hymenoptera: Vespidae)

Summary The frequency of application of ant repellent secreted from the metasomal sternum VI gland to the nest petiole (=rubbing behavior) inParapolybia indica colonies decreased significantly from the preemergence, single-foundress stage to the post-emergence stage. In the pre-emergence stage, rubbing occurred in close temporal association with the departure from the nest, which may maximize the antrepelling effect of the rubbing substance during the period when the nest is not attended by any wasps. On the other hand, only a small proportion of the departures was associated with rubbing in the post-emergence stage.Unlike in other independent-founding polistine wasps, rubbing inP. indica was not associated with petiole enlargement. Among independent-founding polistines,P. indica is unusual in that females enlarge the nest petiole with a large amount of plant fibers instead of strict use of adult saliva, and in that the petiole surface is rough, and may be much more absorbent than that of the secretion petiole. On the other hand, in the species showing a temporal association between petiole enlargement and rubbing, females typically rub immediately after petiole licking (application of oral secretion). This fact, and the lack of such a temporal association inP. indica, may suggest a possibility that the mixture of oral secretion and rubbing substance on the smooth surface of the secretion petiole makes the chemical barrier last longer.     

Host-searching behavior ofVenturia canescens (Grav.) (Hymenoptera: Ichneumonidae): Superparasitism

The foraging behavior ofVenturia canescens, a solitary endoparasitoid of lepidopteran larvae, was investigated in the laboratory. FemaleVenturia canescens with a larger number of mature eggs to lay were found to have higher levels of superparasitism (measured as numbers of eggs laid per parasitized host). Increased parasitoid density was found to result in reduced levels of superparasitism by host-deprived (i.e.,undepleted) wasps. Females which had been allowed access to hosts before the experiment (depleted wasps) laid fewer eggs per parasitized host than undepleted wasps, although there was no significant difference in the levels of superparasitism among the depletion periods of 1, 2, 5, and 7 h. It was also found that an egg which was encountered less than 15 min after oviposition was much less likely to be avoided than one which was encountered after more than 15 min had elapsed.     

Immigration and formation of mixed colonies in red wood ants (Hymenoptera,Formicidae)

Long-term field studies of the composition and spatial structure of settlements of ants of the Formica rufa group were carried out in two regions of Russia (Moscow and Arkhangelsk provinces). Fragmentation of damaged nests followed by reintegration of the fragments is the main way of formation of mixed colonies of ants from different nests (including different species). The principal factor of nest fragmentation is their damage by wild boars, bears, and in some localities, by poachers. The formation of mixed nests and nest complexes with participation of different Formica species was observed. They are formed by joining the ants from several damaged nests or by a colony from a destroyed nest immigrating into an intact one. Regular damage of many nests leads to the formation of broad zones of mixed colonies. The mixed colonies including 2–3 species of wood ants have recently become common. The phenomenon of mixed colonies raises a question as to the relative importance of two basic principles (sociality and specific identity) in the life of ant societies and demonstrates the priority of the social principle.     

Pedestal hairs of the ant Echinopla melanarctos (Hymenoptera, Formicidae): morphology and functional aspects

The South East Asian arboreal Formicine Echinopla melanarctos, as well as some other members of this genus possess a cuticular structure unique in ants, the pedestal hairs. In E. melanarctos, about 700 pedestal hairs are situated on the dorsal and lateral surfaces of the head, the alitrunk, the petiole and the gaster. They are arranged in a polygon-like figuration. On the summit of each of the up to 200-μm high pedestals, a single central hair inserts. This hair (up to 500-μm long) is innervated by a single bipolar mechanosensitive sensory cell. The lumen of each tube-like pedestal contains (1) epithelial cells (2) the sensory cell and the auxiliary cells of the central hair and (3) the long efferent ductules of up to ten isolated bicellular glandular units. Each glandular unit is composed of a secretory glandular cell and a duct cell, all of which are located at the base of a pedestal. The cytoplasm of a glandular cell contains a well-developed end apparatus and is characterised by stacks of smooth and granular endoplasmic reticulum, numerous polyribosomes, a lot of mitochondria and some up to 5-μm large secretory vesicles. The secretion of the gland cells is released on the apex of the pedestal wall via small pores. Approximately 30 μm below their summit, some pedestals possess additionally (up to six) mechanosensitive hairs that are arranged ray-like. We suppose that the pedestal hairs are important in nest-space protection and find that only in ants with high pedestals on the head (Echinopla melanarctos and Echinopla pallipes), the compound eyes are stalked thus overtopping the pedestals.     

Principles of organization of polistinae (Hymenoptera,Vespidae) population

The paper describes some invariant relations of the Polistinae population structure, including resistance to abiotic and biotic factors that occurs against the background of the hierarchy of biological systems and increasing autonomy of their functioning. A decrease in the dependence on the hostile environment is shown to be due to the activity of foundresses and workers adjusting to external rhythms, developing specialized responses to predators and parasites (predictable external noise of biotic nature), and creating new information. The population organization of Polistinae wasps is considered in the framework of Anokhin’s theory of functional systems and systemogenesis. There are specific processes in the population that unite individual colonies and their reproduction; they are accompanied by the formation of an advanced feedback and functional systems. Systemic processes can be simultaneously regarded as “adaptation” (reflecting the organization of environmental elements) and as “adaptiveness” (reflecting the organization of the activity of intra-colony processes and the organization of reproduction). The organization of the colony activity and reproduction in functional systems reflects the future survival rather than the preceding phenomena and events. The behavior of individuals in a colony is determined not only by the effects of abiotic and biotic factors (via transformation of cues into behavioral programs), but also by previous adaptations (stored in the “memory” as images of still absent events). General progress, limited or partial progress, and narrow specialization in the organization of polistine colonies and populations are considered using the examples of morphofunctional, environmental, energy and information criteria. The emphasis on invariant relations makes it possible to more fully describe biological systems in terms of such general categories as isomorphism, homeostasis or self-organization, and also enables us to use more effectively the theory of general functional systems in studying social insects.     

The Role of Passive Defense (Selfish Herd and Dilution Effect) in the Gregarious Nesting of Liostenogaster Wasps (Vespidae, Hymenoptera, Stenogastrinae)

The advantages of the selfish herd and encounter-dilution effects were studied in two congeneric gregarious-nesting species of Stenogastrine wasps, Liostenogaster vechti Turillazzi and L. flavolineata (Cameron), in Malaysia. Results show that nesting in clusters affords these wasps some degree of protection against their main invertebrate predators, parasites and parasitoids. This advantage is reflected in the geometry of the aggregation as well as in the behavior of the individuals. The costs and benefits of group living and the implications of the selection of this nesting strategy in the evolution of sociality in these wasps are discussed.     

Venom induces alarm behaviour in the social wasp Polybioides raphigastra (Hymenoptera: Vespidae): an investigation of alarm behaviour, venom volatiles and sting autotomy

Chemicals from the venom gland elicited alarm behaviour and attack in the Asian polistine wasp Polybioides raphigastra. When presented with crushed venom glands workers of this wasp respond with a mass stinging attack. Gas chromatography–mass spectrometry analyses show that the major volatiles in the venom gland are alkanes, monounsaturated alkenes and 2-alcohols. Several pyrazines, a spiroacetal and aromatics were also identified as trace compounds. The anatomy and morphology of the sting apparatus are reported, and we describe sting autotomy in this wasp. This is the first such report for the Ropalidiinae. The structures responsible for autotomy are likely to be large barbs present on the sting lancets, and a conspicuous tooth present on the medial side of the left lancet. Sting autotomy in P. raphigastra probably plays an important role in the localization of sites of attack by wasps defending the nest.     

Demography of the Okinawan eusocial wasp Ropalidia fasciata (Hymenoptera: Vespidae) I. Survival rate of individuals and colonies, and yearly fluctuations in colony density

We conducted a series of demographic studies of the primitively eusocial wasp Ropalidia fasciata in Okinawa, a subtropical part of Japan, and found the following. The colony cycle of this wasp is annual, spanning from April to November or even December; this is longer than the colony cycles of other temperate polistine wasps so far reported. The survival rate of the marked foundresses was 40% to June, dropping to 1.4% by September. These survival rates are similar to other subtropical and tropical species. Most females that emerged in November were probably second generation adults (progeny of the original foundresses), which would participate in founding nests in the following spring. These facts indicate that R. fasciata in Okinawa is at least partially bivoltine. Survival of a nest to September was 10–20%; however, because a failed nest is often rebuilt, survival of the colony to September was as high as 50%. The mean number of new foundresses produced per foundress was 7.5, and their overwintering survival was 16%. Hence, a single foundress produced, on average, 1.2 progeny foundresses to the following year. Density dependence was shown in the rate at which the progeny foundresses were produced. These results explain the remarkable stability of nest densities from year to year in the area. The above results reveal that R. fasciata in Okinawa shares many demographic characteristics with other primitively eusocial wasps, particularly year‐to‐year stability of nest density and a long colony cycle.     

Chromosome numbers are not fixed in Agaonidae (Hymenoptera: Chalcidoidea)

Agaonidae (the pollinators of fig trees, Ficus spp., Moraceae) are a distinctive family of chalcid wasps with uncertain affinities within the Chalcidoidea. Chromosome numbers have only been described previously for one species (Blastophaga psenes, 2n = 12, the pollinator of F. carica, of subgenus Ficus). In this paper, we used a modified technique to analyze the karyology of three Ceratosolen species, Ceratosolen solmsi, C. graveli and C. emarginatus which are associated with figs of Ficus subgenus Sycomorus. Their karyotypes are 2n = 10. All the chromosomes are metacentric (NF = 20) with little difference in relative lengths, making them hard to distinguish. The variation in chromosome numbers that we have detected suggests that karyology can usefully complement molecular-based studies of the phylogeny of fig wasps.     

Behavior of red wood ants (Hymenoptera,Formicidae) during interaction with different symbiont partners

A comparative analysis of the behavior of Formica polyctena Först during interaction with different symbionts (free-living aphids Aphis grossulariae Kalt. and hidden larvae of the sawfly Blasticotoma filiceti Klug) was carried out. Red wood ants demonstrate different levels of functional differentiation in relatively constant groups of foragers collecting honeydew. A deep “professional” specialization with clear division of a number of tasks among foragers was studied in groups of ants tending aphids. Four professional groups of foragers with different tasks were revealed: “shepherds,” “guards,” “transporters,” and “scouts” (or “coordinators”). The groups of foragers caring for sawfly larvae mainly consist of unspecialized ants. Only few ants (about 5%) remain on duty on the fern plant near B. filiceti larvae and protect the food resource from competitors, especially from other ants. In addition, the ants demonstrate simpler behavior while collecting the larval excretion, resembling that at the sugar feeders. On the whole, the behavior of red wood ants is rather flexible. The level of functional differentiation in groups of foragers collecting honeydew is determined not only by the colony size and requirements but by the nature of their interaction with trophobionts, particularly, by the possibility of direct contact.