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In primitively eusocial societies, all individuals can potentially reproduce independently. The key fact that we focus on in this paper is that individuals in such societies instead often queue to inherit breeding positions. Queuing leads to systematic differences in expected future fitness. We first discuss the implications this has for variation in behaviour. For example, because helpers nearer to the front of the queue have more to lose, they should work less hard to rear the dominant's offspring. However, higher rankers may be more aggressive than low rankers, even if they risk injury in the process, if aggression functions to maintain or enhance queue position. Second, we discuss how queuing rules may be enforced through hidden threats that rarely have to be carried out. In fishes, rule breakers face the threat of eviction from the group. In contrast, subordinate paper wasps are not injured or evicted during escalated challenges against the dominant, perhaps because they are more valuable to the dominant. We discuss evidence that paper-wasp dominants avoid escalated conflicts by ceding reproduction to subordinates. Queuing rules appear usually to be enforced by individuals adjacent in the queue rather than by dominants. Further manipulative studies are required to reveal mechanisms underlying queue stability and to elucidate what determines queue position in the first place. [ABSTRACT FROM AUTHOR]

Abstract: Adult workers of some social insect species show dramatic behavioral changes as they pass through a sequence of task specializations. In the paper wasp, Polybia aequatorialis, female workers begin adult life within the nest tending brood, progress to maintaining and defending the nest exterior, and ultimately leave the nest to forage. The mushroom body (MB) calyx neuropil increases in volume as workers progress from in-nest to foraging tasks. In other social Hymenoptera (bees and ants), MB Kenyon cell dendrites, axons and synapses change with the transition to foraging, but these neuronal effects had not been studied in wasps. Furthermore, the on-nest worker of Polybia wasps, an intermediate task specialization not identified in bees or ants, provides the opportunity to study pre-foraging worker class transitions. We asked whether Kenyon cell dendritic arborization varies with the task specialization of Polybia workers observed in the field near Monteverde, Costa Rica. Golgi-impregnated arbors in the lip and collar calyces, which receive a predominance of olfactory and visual input, respectively, were quantified using Sholl’s concentric circles and a novel application of virtual spherical probes. Arbors of the lip varied in a manner reminiscent of honeybees, with foragers having the largest and in-nest workers having the smallest arbors. In contrast, arbors of the collar were largest in foragers but smallest in on-nest workers. Thus, progression through task specializations in P. aequatorialis involves subregion specific dendritic growth and regression in the MB neuropil. These results may reflect the sensitivity of Kenyon cell dendritic structure to specialization dependent social and sensory experience. [Copyright &y& Elsevier]

The cognitive challenges that social animals face depend on species differences in social organization and may affect mosaic brain evolution. We asked whether the relative size of functionally distinct brain regions corresponds to species differences in social behaviour among paper wasps (Hymenoptera: Vespidae). We measured the volumes of targeted brain regions in eight species of paper wasps. We found species variation in functionally distinct brain regions, which was especially strong in queens. Queens from species with open-comb nests had larger central processing regions dedicated to vision (mushroom body (MB) calyx collars) than those with enclosed nests. Queens from advanced eusocial species (swarm founders), who rely on pheromones in several contexts, had larger antennal lobes than primitively eusocial independent founders. Queens from species with morphologically distinct castes had augmented central processing regions dedicated to antennal input (MB lips) relative to caste monomorphic species. Intraspecific caste differences also varied with mode of colony founding. Independent-founding queens had larger MB collars than their workers. Conversely, workers in swarm-founding species with decentralized colony regulation had larger MB calyx collars and optic lobes than their queens. Our results suggest that brain organization is affected by evolutionary transitions in social interactions and is related to the environmental stimuli group members face. [ABSTRACT FROM AUTHOR]

This study presents the first observations of early-season colony development of Ropalidia plebeiana in Canberra, Australia. The growth pattern of three R. plebeiana nests was measured during weekly observations from October 2006 to January 2007 and showed that nests steadily increase in size over the early summer to approximately 50 cells when the nest is newly established and to approximately 170 cells when nests from the previous season are re-used. A first generation of adult females is produced by December, and the bimodality of the curves of egg, larval and pupal numbers indicates that these three developmental stages last approximately 2–3 weeks each. The nesting cycle of R. plebeiana in Canberra commences approximately 2 weeks later than in coastal Australia, the shorter summers in this inland region restricting the length of the active season. R. plebeiana did not form dense nesting aggregations in Canberra as described elsewhere, with only small colonies consisting of a single or just a few nests. Characteristic comb-cutting behavior of the species was observed but this did not result in complete nest division as recorded from coastal populations. [ABSTRACT FROM AUTHOR]

We analysed morphological variation among 17 forewing characters within five populations of the paper wasp, Polistes dominulus, in Iran. The raw planar coordinate data were aligned using geometric and mathematical calculations in kendall's shape space. After transfer of the data to a linear euclidean space, i.e., tangent space, multivariate analysis of 135 images of forewings were made using their geometric morphometric characters (30 in the forewings). We observed a direct correlation between morphological characters and the geographically easiest travel distance along river valleys and mountain ranges. [ABSTRACT FROM AUTHOR]

We provide the first description of the mating system, genetic structuring and dispersal in the Australian paper wasp, Polistes humilis. Individuals were collected from 13 colonies that were within 700m of each other at a location near Sydney, Australia. Analysis of genotypic data from four microsatellite loci provided no evidence of males siring offspring in their natal colony and heterozygote excesses within most colonies suggest that this form of outbreeding is typical in P. humilis. The same data show that queens are singly mated and that the number of queens and therefore groups of full siblings increase with colony size. Consequently, genetic variation also increases with colony size. The relationship between the number of queens and colony size in P. humilismay be indicative of a breakdown in dominance hierarchies in larger colonies or a defence against disease transmission. [ABSTRACT FROM AUTHOR]

Abstract: Stings by Polistes wasps can cause life-threatening allergic reactions, pain and inflammation. We examined the changes in microvascular permeability and neutrophil influx caused by the venom of Polistes lanio a paper wasp found in southeastern Brazil. The intradermal injection of wasp venom caused long-lasting paw oedema and dose-dependently increased microvascular permeability in mouse dorsal skin. SR140333, an NK1 receptor antagonist, markedly inhibited the response, but the NK2 receptor antagonist SR48968 was ineffective. The oedema was reduced in capsaicin-treated rats, indicating a direct activation of sensory fibres. Dialysis of the venom partially reduced the oedema and the remaining response was further inhibited by SR140333. Mass spectrometric analysis of the venom revealed two peptides (QPPTPPEHRFPGLM and ASEPTALGLPRIFPGLM) with sequence similarities to the C-terminal region of tachykinin-like peptides found in Phoneutria nigriventer spider venom and vertebrates. Wasp venom failed to release histamine from mast cells in vitro and spectrofluorometric assay of the venom revealed a negligible content of histamine in the usual dose of P. l. lanio venom (1nmol of histamine/7μg of venom) that was removed by dialysis. The histamine H1 receptor antagonist pyrilamine, but not bradykinin B1 or B2 receptor antagonists, inhibited venom-induced oedema. In conclusion, P. l. lanio venom induces potent oedema and increases vascular permeability in mice, primarily through activation of tachykinin NK1 receptors by substance P released from sensory C fibres, which in turn releases histamine from dermal mast cells. This is the first description of a neurovascular mechanism for P. l. lanio venom-mediated inflammation. The extent to which the two tachykinin-like peptides identified here contribute to this neurogenic inflammatory response remains to be elucidated. [Copyright &y& Elsevier]

The establishment of the multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), in North America has resulted in negative impacts on fruit production. We investigated the overwintering survival of H. axyridis after feeding on four diets: injured wine grape berries, 25% sucrose solution, water, and a control containing no food or water. After being exposed to these diets for 6 days, live individuals were transferred to clean plastic Petri dishes, and held at 5 ± 1 °C in growth chambers throughout the winter. Survival was recorded every month. Adult lady beetles collected during the overwintering flight in mid-October had higher survival rates than beetles collected from soybean fields in mid-August. These results suggest that an adaptation period prior to diapause increases the chances of lady beetle survival over the winter. In addition, injured wine grape berries, sugar, and water decreased beetle mortality during the overwintering months. Our results also showed that under similar conditions, females have higher survival during the winter than males. The importance of sugar and water on winter survival may drive H. axyridis adults to vineyards for feeding on wine grapes. Finally, we tested if adults of H. axyridis and the European paper wasp, Polistes dominulus Christ (Hymenoptera: Vespidae), were able to break the grape skin. Harmonia axyridis adults and paper wasps were not able to cause the primary injury to berries of Frontenac grapes under laboratory conditions. These results suggest that control of paper wasps in vineyards may not affect H. axyridis aggregations, and that H. axyridis feeding on wine grapes depends on previous injury to grape berries. [ABSTRACT FROM AUTHOR]

In animals, restlessness is a strategy to fulfil a goal within a narrow seasonal window when ecological and physiological constraints require the performance of certain fundamental events. Although restlessness can be expected in parasitic species that have their biological cycle similar to that of the host, no ‘urge to parasitize’ has thus far been demonstrated in any host-parasite pair. The paper wasp, Polistes sulcifer, is a social parasite that has lost the nest-building ability and marked by the absence of the worker caste; it depends on a congeneric host species to reproduce. Host colony usurpation is successful when it occurs in a well-defined seasonal window. The short time available for host-nest usurpation might lead to parasite restlessness that overlaps this period. Under controlled laboratory conditions, we demonstrated that the parasite females exhibit ‘usurpation restlessness’ during the period in which field usurpations occur. Additional experiments showed that both this period and laboratory temperature influence the parasite’s ‘urge to usurp’. [ABSTRACT FROM AUTHOR]

Foundresses and workers of Polistes paper wasps show slight morphological and physiological differences. However, after the emergence of the workers, the castes can be readily discriminated by their behaviour: the dominant foundress is the principal egg-layer, whereas workers perform different tasks linked to colony development. Previous studies have demonstrated in this genus that defence of the colony by the workers is more effectively carried out by a collective response elicited by venom volatiles used as alarm pheromones. In the present study, gas chromatography-mass spectrometry analyses of the venom volatiles of foundresses and workers of Polistes dominulus (Christ) show predominantly quantitative differences. Spiroacetals, mainly (E,E)-2,8-dimethyl-1,7-dioxaspiro[5.5]undecane, are significantly higher in the venom volatiles fraction of workers, whereas the amount of N-(3-methylbutyl)acetamide is almost double in foundresses. On the basis of the chemical results, behavioural assays were performed on fifteen field colonies to test the alarm response of the resident wasps to venom extracts from foundresses and workers. Our behavioural results suggest that worker venom has a stronger alarm effect on the colonies than that of the foundresses, which seems unable to elicit the complete alarm response ending with a final attack and sting. The venom volatiles of P. dominulus workers serve mainly to alarm the colony whilst those of foundresses may also be linked to additional functions related to conspecific interactions. [ABSTRACT FROM AUTHOR]

Indirect benefits obtained through the reproduction of relatives are fundamental in the formation and maintenance of groups. Here, we examine the hypothesis that females of the temperate paper wasp Polistes dominulus preferentially form groups with close relatives. Genetic relatedness data were obtained for 180 groups of females collected at the early stages of the nesting cycle of a large population of P. dominulus in two sites in southwestern Spain. Average within-group relatedness values ranged from 0.189 to 0.491. Foundresses on early nests were significantly more closely related than females in winter aggregations or in stable groups (just before workers emerged). Within-group relatedness values were independent of group size. The vast majority of worker-producing nests ( c. 85%) had one or more females that were unrelated (or distantly related) to the remaining members of the group. These results provide further support to the hypothesis that indirect fitness benefits alone are unlikely to explain why P. dominulus foundresses form cooperative associations. [ABSTRACT FROM AUTHOR]

To establish a dominance order, social animals often rely on indicators of fighting to avoid costly aggressive encounters. In some species, individuals use colour patterns to signal their social status. Recent studies claimed that facial markings in the eusocial paper wasp Polistes dominulus are status badges that allow co-foundresses to form a linear hierarchy based on individual quality. Here, we evaluated facial patterns in natural populations of P. dominulus, in its native range, to observe whether the marks reflect overall wasp quality in different contexts. We used the same measures of clypeus patterns used by earlier studies, but did not find that they functioned as status badges. Our analyses showed no evidence that visual markers are related to: (i) size, (ii) probability of surviving winter, (iii) social rank in spring associations, or (iv) health status (assessed by the presence of strepsipteran endoparasites). Size, however, is important. Larger wasps are more likely to survive the winter and to acquire the dominant position in spring associations. Larvae infected with endoparasites become smaller adult wasps. These findings suggest that body size is a reliable quality indicator on which wasps build their social networks, and that clypeus patterning is not involved. [ABSTRACT FROM AUTHOR]

Abstract  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. [ABSTRACT FROM AUTHOR]

Abstract  In temperate climates, female paper wasps typically initiate new colonies in the spring. Several nest-founding tactics have been documented in Polistes species, including solitary nest initiation, joining a cooperative association, usurping an existing nest, or adopting an abandoned nest. Occasionally, exceptionally large groups of females have also been found reusing nests from the previous season. Here we report this phenomenon in introduced populations of the Eurasian species Polistes dominulus. We describe in detail the demographic and genetic characteristics of one such spring colony from Los Angeles, California, USA, which was collected with 84 associated adults and all stages of developing brood in its 613 cells. Genetic and morphological data indicate the presence of multiple reproductively active females of varying relatedness, as well as many nonbreeding females, including probable early-produced offspring. Despite some evidence of chaotic social conditions, the colony appeared to have been highly productive. Additional observations of similar colonies are needed to determine how control is maintained within such a large breeding aggregation. [ABSTRACT FROM AUTHOR]

Using demography data for the primitively eusocial wasp Ropalidia fasciata collected at 10 survey stations over 5 years, the effects of foundress group size (FGS) on colony survival rate and production of progeny were examined. The distribution pattern for the frequency of nests established by different numbers of foundresses fit a 0-truncated negative binomial distribution. The rate of nest failure up to the beginning of June decreased with FGS, and colony survival rate up to June and September increased with FGS. Although there were large variations among stations and years, the survival rate of colonies established by a single foundress (haplometrotic colonies) was significantly lower than that of colonies established by two or more foundresses (pleometrotic colonies). The number of new adults that emerged per colony up to the end of July increased with FGS, but there was no significant correlation between number of new adults per foundress and FGS. The number of potential foundresses produced per colony tended to increase with FGS, but there was no significant difference among the values produced per foundress for the three FGS categories. The percentage parasitism by an ichneumon parasitoid, Arthula formosana, tended to be higher in colonies established by a small number of foundresses. The relation between FGS and the productivity of the new foundresses was not statistically significant, suggesting that independent founding may be a better strategy for future subordinate foundresses. However, the long colony life span (mean 100 days, maximum 240 days) as compared with the shorter life span of foundresses (40 or 50 days) may be a good condition for the coexistence of many egg-layers in a colony. [ABSTRACT FROM AUTHOR]

Abstract: Several studies have shown that differences in the relative abundance of cuticular hydrocarbons occur between reproductives and non-reproductives in many social insects. These differences also exist between Polistes dominulus foundresses and their first emerged daughters (usually indicated as workers), but they gradually disappear when workers from orphaned colonies develop their ovaries and assume a reproductive role in the colony. However, hydrocarbons are not the exclusive components of cuticular layer of wasps. Mass-spectrometry analysis of cuticular methanol extracts from Polistes paper wasps showed a complex pattern of polar substances, partly or totally proteinaceous in nature. We found that these compounds, ranging from 918 to 2679Da, showed a clear caste differentiation between foundresses and their first emerged daughters (usually indicated as “workers”), both in queen-right and in orphaned colonies. Conversely to hydrocarbons, workers from orphaned colonies maintain a significant difference from foundresses in the pattern of the medium molecular weight (MW) polar compounds obtained by MALDI-TOF. On the basis of such results we hypothesize that a reliable cue to identify foundresses from daughters, and not only their reproductive status, may exist in Polistes wasps. Although the great majority of previous work on social insect communication has focused on cuticular hydrocarbons, our findings suggest that the medium MW component of cuticular substances may be involved in recognition. [Copyright &y& Elsevier]

Unique among insects, some paper wasp species recognize conspecific facial patterns during social communication. To evaluate whether specialized brain structures are involved in this task, we measured brain and brain component size in four different paper wasp species, two of which show facial pattern recognition. These behavioral abilities were not reflected by an increase in brain size or an increase in the size of the primary visual centers (medulla, lobula). Instead, wasps showing face recognition abilities had smaller olfactory centers (antennal lobes). Although no single brain compartment explains the wasps’ specialized visual abilities, multi-factorial analysis of the different brain components, particularly the antennal lobe and the mushroom body sub-compartments, clearly separates those species that show facial pattern recognition from those that do not. Thus, there appears to be some neural specialization for visual communication in Polistes. However, the apparent lack of optic lobe specialization suggests that the visual processing capabilities of paper wasps might be preadapted for pattern discrimination and the ability to discriminate facial markings could require relatively small changes in their neuronal substrate. Copyright © 2007 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

1. In insect communities on fermented tree sap, agonistic interactions occur frequently among the component species. This study examined the effects of such interference competition on community structure and species co-occurrence on a spatio-temporally small scale. 2. Experimental removal of giant hornets ( Vespa mandarinia), one of the most dominant species, showed no effect on either species richness or total abundance of the remaining insects. Significant increases in abundance were detected for five out of 19 taxa: V. ducalis, V. crabro, Lasius japonicus, Cryptarcha lewisi, and a group of micro sap beetles ( Epuraea spp., Haptoncus spp., and Haptoncurina spp.). 3. Behavioural observations indicated that interference did not alter the staying time at a patch (an exuding spot on a tree) for V. mandarinia, Rhomborrhina japonica, and Neope goschkevitschii. Interference-mediated departure from a patch was less frequent than departure without interference for all nine species observed, except for R. japonica. These behavioural contexts varied yearly for several species. 4. These results suggest that interference competition is a minor determinant of overall community structure, despite its conspicuousness. This weak effect of interference may be attributed to the short stay of dominant species and to individual behavioural responses, such as competitor avoidance. The variable influence of interference might be related to differences in species attributes, such as dominance hierarchies. [ABSTRACT FROM AUTHOR]

The expression of alternative traits that benefit eusocial individuals but are not directly involved in reproductive differences among those individuals, which I call ‘eusocially selected traits’, may vary in response to environmental changes if this increases an individual's inclusive fitness. In this study, I describe traits that separate individuals within the reproductive division of labor of Mischocyttarus mexicanus, a eusocial paper wasp, and determine whether observed eusocially selected traits vary across seasons. I examined M. mexicanus because females initiate new nests throughout most of the year where they experience different conditions depending on the season. Findings from this study suggest two main conclusions: (1) phenotypic differences among M. mexicanus females are mixed, showing specialized, generalized, and context-dependent eusocially selected traits and (2) a female's position within the reproductive division of labor may be influenced by its state. The presence of context-dependent traits, e.g. large females initiated solitary nests in the spring and grouped nests during the summer, suggests that the payoff for pursuing different positions within the reproductive division of labor changes across seasons. The expression of context-dependent eusocially selected traits also suggests that, roles, instead of castes, may better reflect the reproductive division of labor among individuals of eusocial species like M. mexicanus. [ABSTRACT FROM AUTHOR]

The abdominal wagging behavior of Polistes dominulus foundresses is highly conspicuous and frequently performed, but comprehensive descriptions are lacking. It has been suggested that abdominal wagging is involved in vibratory signaling, but the wagging behavior has not been distinguished from the substrate-borne signal, and nest vibrations have never been described. Here I provide a detailed description of the behavior and the first video-linked accelerometer recordings of nest vibrations. These recordings indicate that an impulsive vibration is produced when the adult female repeatedly strikes its abdomen against cell rims as it swings it across the nest surface. These strong vibrations propagate throughout the nest. Scanning electron microscopy reveals no specialized structures for vibration production on the abdomen or nest. These results provide a foundation for the mechanical playback studies required to experimentally determine the function of abdominal wagging. [ABSTRACT FROM AUTHOR]

The mushroom bodies (MB) are a complex neuropil in insect brains that have been implicated in higher-order information processing such as sensory integration and various types of learning and memory. Eusocial insects are excellent models to test functional neural plasticity in the MB because genetically related nest mates differ in task performance, environmental experience and social interactions. Previous research on eusocial insects shows that experience-dependent changes in brain anatomy (i.e., enlarged MB calyces) are positively correlated with task performance and social interactions. In this study, we quantified relationships of task performance and social and reproductive dominance with MB volume in Polistes instabilis, a primitively eusocial paper wasp. We used experimental removals of dominant workers to induce changes in aggressive behavior and foraging by workers. Ovary development and social dominance were positively associated with the volume of the MB calyces relative to the region containing the Kenyon cell bodies. In contrast to highly eusocial insect workers, foraging behavior was not positively correlated with MB calycal volume. We conclude that mushroom body volume is more strongly associated with dominance rank than with foraging behavior in Polistes instabilis. Copyright © 2007 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

Given the diversity of animal signals, there has been recent interest in categorizing signals into probable functions according to their properties. For example, models predict that signals of quality should be costly and condition dependent, whereas signals of individual identity should be cheap and expressed independently of condition. Here, we test these predictions by comparing the condition dependence of signals of individual identity and quality in Polistes wasps. Polistes fuscatus wasps have black and yellow patterns on the face and abdomen that signal individual identity, whereas Polistes dominulus wasps have black and yellow facial patterns that signal aspects of quality related to dominance. We reared both species with and without supplemental food and examined the facial patterns of the resulting offspring. As predicted, food availability did not influence the development of identity signals in P. fuscatus. In strong contrast, P. dominulus wasps reared with supplemental food had facial patterns that signaled higher levels of quality than P. dominulus reared without supplemental food. Interestingly, the identity and quality signals have different condition dependence, despite being composed of similar pigments, suggesting that signal function has a stronger influence on signal properties than pigmentation. Because body size is often correlated with quality signal elaboration, we also tested how food supplementation influenced offspring size. In both species, supplemented colonies produced smaller offspring than nonsupplemented colonies, suggesting that queens may invest in producing fewer, larger offspring in stressful environments. [ABSTRACT FROM PUBLISHER]

Animals that live in cooperative societies form hierarchies in which dominant individuals reap disproportionate benefits from group cooperation. The stability of these societies requires subordinates to accept their inferior status rather than engage in escalated conflict with dominants over rank. Applying the logic of animal contests to these cases predicts that escalated conflict is more likely where subordinates are reproductively suppressed, where group productivity is high, relatedness is low, and where subordinates are relatively strong. We tested these four predictions in the field on co-foundress associations of the paper wasp Polistes dominulus by inducing contests over dominance rank experimentally. Subordinates with lower levels of ovarian development, and those in larger, more productive groups, were more likely to escalate in conflict with their dominant, as predicted. Neither genetic relatedness nor relative body size had significant effects on the probability of escalation. The original dominant emerged as the winner in all except one escalated contest. The results provide the first evidence that reproductive suppression of subordinates increases the threat of escalated conflict, and hence that reproductive sharing can promote stability of the dominant–subordinate relationship. [ABSTRACT FROM AUTHOR]

During the past 15 years in the Ina Valley, Japan, mature fifth instar larvae of the yellow hornet, Vespa simillima, have been occasionally ejected from colonies during September. During 2005, this unusual behavior was particularly widespread, and collection of several V. simillima colonies confirmed that very few fifth instar larvae were present. When compared with an average colony, constructed from 41 colonies collected 20 years previously, colonies in 2005 had 80% fewer fifth instar larvae, despite queen egg-laying rates remaining similar. It may be that the amount of food provided to the larvae by workers is less than the amount of larval secretions received by workers, which is causing the larvae to become emaciated and preventing them from pupating. This phenomenon normally occurs naturally at the end of the colony cycle in November, when prey abundance decreases and larval secretions or other sources of carbohydrates, for example from ripe fruit, tree sap, or aphid secretions, are needed by sexuals to build up fat reserves. However, another possibility is that the unexplained appearance of this phenomenon in September, which is causing colonies to collapse without producing any or very few sexuals, is due to foragers feeding the larvae prey that are contaminated with insect growth regulators, via pesticides, which are known to prevent successful pupation. [ABSTRACT FROM AUTHOR]

Only two out of 959 pre-emergence colonies of the paper wasp Polistes snelleni de Saussure surveyed between 1989 and 1996 in Sapporo, northern Japan, were found to be two-foundress colonies, and the others were single-foundress colonies. The two foundresses in one of the two colonies showed neither aggressive dominance behavior nor clear division of labor between them during a total of approximately 30 h in the first half of the pre-emergence stage. Although both of the foundresses foraged for pulp and laid eggs, only one foundress foraged for prey and delivered it to the other. [ABSTRACT FROM AUTHOR]

The idea that insects play has often aroused skepticism. Nevertheless, the authors investigated the occurrence of a playlike behavior in young individuals of a paper wasp. Polistes dominulus foundresses hibernate in aggregations and found associative nests in the spring. In aggregations, wasps exhibit most of the rank-dependent behaviors that they will use in the nest 6 months later. In aggregations, precocious dominance interactions are performed with exaggerated frequencies, whereas aggression and trophallaxis are rare. Dominance behavior in aggregations is apparently useless, but it probably allows wasps to assess their dominance potential without any apparent reproductive competition. Surprisingly, these interactions may be best interpreted as play because dominance behavior in aggregations shares several features with mammalian play fighting. [ABSTRACT FROM AUTHOR]

The active and coordinating capacity of defending the nest is a key feature of social insects. The present study investigates the presence of alarm pheromones in the venom of workers of the social wasp, Polistes dominulus. Laboratory experiments were performed with caged colonies of P. dominulus using a wind tunnel apparatus to test the behavioural response of workers to venom released by other workers and to venom extracts. Contrary to that previously reported for European paper wasps, the present results show that the venom is the source of alarm pheromones. Field experiments combining a visual (black target) and a chemical stimulus (venom extract) were performed to test the effect of the venom on the reaction of colonies. Wasps leave the nest, land on the visual target and attack the target significantly more once exposed to venom extract plus target than to solvent plus target. This work shows that the venom of P. dominulus workers elicits an alarm response, reduces the threshold for attack and acts as an attractant on targets. These results using P. dominulus indicate that, in both American and European species, colony defence is based on the same features, suggesting that chemical alarm is a widespread trait in the genus Polistes. [ABSTRACT FROM AUTHOR]

Ropalidia marginata and Ropalidia cyathiformis are two Old World, primitively eusocial, tropical polistine wasps that exhibit perennial, aseasonal, indeterminate nesting cycles. Queens are periodically ousted and replaced by one of the workers, whom we refer to as the potential queen. Here we identify the characters of the potential queens by experimentally removing queens from several colonies of both species. Potential queens in R. marginata are unspecialized, worker-like individuals, not unique in their dominance ranks. In contrast, potential queens in R. cyathiformis are queenlike individuals and unique in always holding the top dominance rank among the workers. We suggest that this striking difference in the behaviour of the potential queens of the two species has to do with the very different mechanisms that queens of the two species use to suppress worker reproduction. In regard to their ovarian development, potential queens of neither species are unique and they are only one among several individuals with partially developed ovaries. This may have to do with the fact that queen replacements are frequent in tropical, aseasonal climates, making it adaptive for several individuals to be prepared to take over the position of the queen at short notice. [ABSTRACT FROM AUTHOR]

Abstract: The wasps of the genus Polistes (paper wasps), have a worldwide distribution and are widespread in human-built areas. Like other social wasps, they are at the top of food chains and are therefore exposed to the dangers of biomagnification, given that the larvae are fed predominantly with prey that consist of herbivorous insects. The larval faeces, larval fecal masses, in the form of a semi-solid ball, are made up of the residues of the diet of the larva, which are emitted and compressed on the floor of the cell during the larval metamorphosis. Larval fecal masses may accumulate lead (up to 36 times with respect to the adult body), therefore they were used as substrate for the analysis. From the analysis of sample nests of Polistes dominulus in various sites of the urban area of Florence, it emerges that the larval fecal masses are an analytical substrate with which it is possible to distinguish zones with differing degrees of lead pollution. The lead concentration measured in the larval fecal masses turns out to be directly correlated with vehicle traffic density, the main lead source in Florence when the survey was carried out. The notable increase in the lead concentration of larval fecal masses from the rural to the urban nest (11.15 times), in contrast with the much more limited level of pupae (4.39 times), seems to indicate the efficiency of the excretion and/or barrier mechanisms. These wasps seem to be a promising species for biomonitoring lead pollution in order to better understand its dynamics in anthropic ecosystems after the progressive diffusion of unleaded gasoline. [Copyright &y& Elsevier]

Female Polistes paper wasps can initiate colonies either solitarily or in cooperative groups. Reproduction is often distributed unequally in groups, even to the point of complete monopolization of breeding by the dominant group member. Transactional models of reproductive skew predict the degree of reproductive partitioning, assuming that the dominant controls group membership and will yield a proportion of reproduction to a subordinate as an incentive to stay peacefully in the group. Using a combination of demographic, genetic and morphological data from a population of P. dominulus, we test predictions of ‘classical’ two-person skew models as well as more complex ‘N-person’ models. This is the most comprehensive study of skew in this species to date, and the results generally do not support transactional models. We found no relationship between skew and relatedness for dyads, and complete skew was observed in unrelated groups despite the prediction for this population that such groups should not occur. In contrast to N-person model predictions, group size tended to increase with relatedness. Although we did find the predicted positive correlation between group size and skew for groups of nonrelatives, this relationship was weak. The zone of conflict between the predicted minimum and maximum staying incentives often spans the entire possible range of reproductive skew, suggesting that a ‘tug-of-war’ scenario may be more appropriate than a transactional framework for understanding within-colony dynamics. Overall, our results demonstrate that transactional skew models have little predictive power and are therefore unlikely to yield further insight into Polistes wasp societies. [Copyright &y& Elsevier]

Nineteen species of the paper wasp genus Ropalidia are recorded from Vietnam, of which three are new records for Vietnam. A key to all of the species, distribution records and taxonomic remarks are given. New synonymies proposed are: Ropalidia bicolorata parvula van der Vecht, 1962, under R. bicolorata van der Vecht, 1962; R. hongkongensis juncta van der Vecht, 1941, under R. hongkongensis (de Saussure, 1854); Icaria jucunda Cameron, 1898, and R. marginata sundaica van der Vecht, 1941, under R. marginata (Lepeletier, 1836); R. rufocollaris atrata van der Vecht, 1941, under R. rufocollaris (Cameron, 1900); and R. stigma rufa van der Vecht, 1941 and R. stigma nigrolineata van der Vecht, 1962, under R. stigma (Smith, 1858). [ABSTRACT FROM AUTHOR]

Abstract: Ropalidia marginata is a primitively eusocial, polistine wasp widely distributed in peninsular India. In spite of its primitively eusocial status, queens of R. marginata are surprisingly docile and behaviourally non-dominant (except during the first week or so of their careers as queens). Yet they successfully maintain reproductive monopoly throughout their careers, probably through the use of pheromones. Workers exhibit dominance–subordinate interactions but these behaviours are not involved in regulating reproductive competition among the workers because workers with high dominance ranks are not necessarily the ones who replace lost queens. We have speculated and provided correlational evidence before that dominance-subordinate interactions among the workers have been co-opted in this species for the workers to regulate each other''s foraging. Here, we provide experimental evidence in support of the speculation, by reducing demand for food and showing that this results in a significant decrease in the frequency of dominance-subordinate interactions among the workers. [Copyright &y& Elsevier]

A striking feature of hymenopteran societies is the general absence of male workers. There are some reports of males feeding larvae in some social wasps, but not in the extensively studied Ropalidia marginata. By providing excess food and by removing the females, we investigated whether males are incapable of feeding larvae or lack opportunities to do so. Males fed larvae with probabilities and rates comparable to those of females, suggesting that lack of preadaptation to feed larvae does not explain the absence of male workers in social Hymenoptera. Although males were not as efficient as females at feeding larvae, they seemed capable of doing enough for natural selection to have promoted the evolution of male workers if there were not other factors preventing it. Genetic relatedness asymmetry may be one such factor. A third hypothesis for the absence of male workers in social Hymenoptera, concerning the increased susceptibility of haploid males to infection, is not relevant because males handle and masticate prey for their own consumption and otherwise interact with the larvae. [Copyright &y& Elsevier]

Hyaluronidase (E.C. 3.2.1.35), one of the three major allergens of yellow jacket venom, is a glycoprotein of 45 kDa that is largely responsible for the cross-reactivity of wasp and bee venoms with sera of allergic patients. The asparagine-linked carbohydrate often appears to constitute the common IgE-binding determinant. Using a combination of MALDI MS and HPLC of 2-aminopyridine-labelled glycans, we found core-difucosylated paucimannosidic glycans to be the major species in the 43–45 kDa band of Vespula vulgaris and also in the corresponding bands of venoms from five other wasp species ( V. germanica, V. maculifrons, V. pensylvanica, V. flavopilosa and V. squamosa). Concomitant peptide mapping of the V. vulgaris 43 kDa band identified the known hyaluronidase, Ves v 2 (SwissProt P49370), but only as a minor component. De novo sequencing by tandem MS revealed the predominating peptides to resemble a different, yet homologous, sequence. cDNA cloning retrieved a sequence with 58 and 59% homology to the previously known isoform and to the Dolichovespula maculata and Polistes annularis hyaluronidases. Close homologues of this new, putative hyaluronidase b (Ves v 2b) were also the major isoform in the other wasp venoms. [ABSTRACT FROM AUTHOR]

Abstract: We analysed changes in cuticular hydrocarbon signatures of workers in orphaned colonies of the paper wasp Polistes dominulus. In natural conditions, workers and foundresses possess characteristic cuticular signatures, and foundresses are further distinguishable, both behaviourally and chemically, on the basis of their rank in a reproductive dominance hierarchy. In our study, several workers were found to develop their ovaries and produce cuticular signatures resembling those of dominant foundresses, while remaining workers possessed undeveloped ovaries and had cuticular blends characteristic of subordinate foundresses. Workers that did not develop their ovaries had changed epicuticular signatures, demonstrating that the mixture of hydrocarbons of worker individuals is strongly dependent on social role and environment. Our results suggest that the composition of epicuticular lipids is not determined at the pre-imaginal stage, and that physiological pathways leading to cuticular chemical changes are similar in foundresses and workers of P. dominulus. [Copyright &y& Elsevier]

Assessing a conspecific's potential is often crucial to increase one's fitness, e.g. in female choice, contests with rivals or reproductive conflicts in animal societies. In the latter, helpers benefit from accurately assessing the fertility of the breeder as an indication of inclusive fitness. There is evidence that this can be achieved using chemical correlates of reproductive activity. Here, we show that queen quality can be assessed by directly monitoring her reproductive output. In the paper wasp Polistes dominulus, we mimicked a decrease in queen fertility by regularly removing brood. This triggered ovarian development and egg-laying by many workers, which strongly suggests that brood abundance is a reliable cue of queen quality. Brood abundance can be monitored when workers perform regular brood care in small size societies where each brood element is kept in a separate cell. Our results also show that although the queen was not manipulated, and thus remained healthy and fully fertile, she did not control worker egg-laying. Nevertheless, when workers laid eggs, the queen secured a near reproductive monopoly by selectively destroying these eggs, a mechanism known as ‘queen policing’. By contrast, workers destroyed comparatively few queen-laid eggs, but did destroy each other's eggs. [ABSTRACT FROM AUTHOR]

The taxonomy of the Australian species of twoRopalidiaspecies groups, theR. stigmagroup andR. variegatagroup, is revised and their distribution patterns are discussed. Two species in theR. stigmagroup (R. darwiniandR. elegantula), and four species in theR. variegatagroup (R. flavinoda,R. gregaria,R. mutabilisandR. revolutionalis) are recognized in Australia, with a new synonymy ofR. mutabilis torresianaRichards, 1978, underR. gregaria(de Saussure, 1854).Icaria torridaSmith, 1863, from Seram Island is synonymized withR. unicolor(Smith, 1859). [ABSTRACT FROM AUTHOR]

To learn the evolutionary trajectories of caste differentiation ineusocial species is a major goal of sociobiology. We present anexplanatory framework for caste evolution in the eusocial wasp genusPolistes (Vespidae), which is a model system for insect eusocialevolution. We hypothesize that Polistes worker and gyne castes stem fromtwo developmental pathways that characterized the bivoltine life cycleof a solitary ancestor. Through individual-based simulations, we showthat our mechanistic framework can reproduce colony-levelcharacteristics of Polistes and, thereby, that social castes can emergefrom solitary regulatory pathways. Our explanatory frameworkillustrates, by specific example, a changed perspective forunderstanding insect social evolution. [ABSTRACT FROM AUTHOR]

Mating structure and genetic relatedness among gynes (potential reproductive females) of the paper waspPolistes snelleniwere investigated using DNA microsatellite markers. All colonies had one foundress inseminated by a single male, and no sign of inbreeding was detected. The mean genetic relatedness among gynes was 0.734 ± 0.028 (SE), which is not significantly different from the expected value of 0.75 for full sisters. [ABSTRACT FROM AUTHOR]

We conducted a series of demographic studies of the primitively eusocial waspRopalidia fasciatain 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 thatR. fasciatain 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 thatR. fasciatain Okinawa shares many demographic characteristics with other primitively eusocial wasps, particularly year-to-year stability of nest density and a long colony cycle. [ABSTRACT FROM AUTHOR]

Grooming is a commonly observed behavior in many animals. One function of grooming is to clean the body of debris and parasites. An additional function may be to homogenize chemical cues present on the body. This latter purpose is especially likely in species in which contact-based chemical communication occurs, such as in eusocial insects. In this study we address the context, sequence, frequency and duration of 683 acts of self-grooming performed by the paper wasp,Polistes dominulus. In general, individuals groomed heads after cell inspections, and abdomens after sitting, suggesting that grooming serves to remove debris from the body. Although no differences were observed in the total amount of time spent grooming, foundresses groomed significantly more often than did workers. Wasps were equally likely to groom thoraces or abdomens following heads, but were more likely to groom abdomens after thoraces and heads after abdomens. Interestingly, the appendages used to groom individual body parts were highly specific (e.g. the prothoracic legs were used for the head), thus indicating that grooming is not used to homogenize chemical cues across the body surface of the wasp. [ABSTRACT FROM AUTHOR]

We examined the population genetic structure of an extremely successful invasive wasp, Polistes dominulus. Although successful biological invasions of social insects have been associated with genetic bottlenecks, our research uncovered an unexpected level of genetic diversity in the northeastern U.S. invasion population. Compared with a previously studied European sample, the northeastern U.S. invasion population shows no significant reduction in gene diversity and no trace of a genetic bottleneck in the putative "introduction population." We identified multiple private microsatellite alleles in both Massachusetts and New York, which strongly suggests that the northeastern U.S.P. dominulus population arose from at least two independent introductions. Although a genetic bottleneck may enhance invasion success for some social insects, genetic and geographical data on this successful invader suggest that this wasp may represent the converse. Our results support immediate identification of genetic diversity in an invasion population before the occurrence of secondary introductions as an essential part of managing and controlling invasive species. [ABSTRACT FROM AUTHOR]

This study investigated the effects of alarm pheromone components on the heat production rates of hornets (Vespa crabro) by means of direct calorimetry. In a flow-through system, pheromones from hornets, honeybees (Apis mellifera) and yellowjackets (Vespula vulgaris) were sucked through a measuring vessel containing a group of hornet workers. The locomotive reaction of hornet workers was recorded as an increase of the heat production rate. Hornets exhibited a strong response to their own alarm pheromone components, mainly 2-methyl-3-butene-2-ol (MBO). They also reacted intensively upon the main alarm pheromone component of the honey bee, isopentylactetate (IPA), but less pronounced to alarm pheromone components of yellowjackets. The metabolic response of hornets to MBO was dose-dependent. The heat production rates of provoked hornet workers were similar to those of flying hornets. (z)-9-Pentacosene, a substance which is believed to be a thermoregulative brood pheromone, induced no metabolic reaction. [Copyright &y& Elsevier]

The aim of this paper is to provide the first complete description of ventilating activity undertaken by the Oriental hornet. At the portal of each nest of the Oriental hornet ( Vespa orientalis, Hymenoptera, Vespinae) there are worker hornets that beat their wings in synchrony with one another, during summer months and at the onset of daylight. These ventilating hornets stand at regular intervals around the nest entrance, not touching, and with their heads facing largely outwards. Although the ventilating hornets stand apart from one another, their wings beat at a uniform pace and there is no connection whatsoever between these hornets and the sentinel worker hornet (or hornets) that inspect those entering the nest. The ventilatory wing beating lasts 3.4 min on average, and upon completion, the ventilating worker either emerges to forage in the field or more often retreats back into the nest. Young queens have seldom been seen ventilating and males have never been seen engaged in such activity. Ordinarily, no ventilatory activity takes place upon the brood combs inside, such as is observed in artificial breeding boxes with a transparent wall, but on rare occasions ventilation may also occur on the inner side of the portal. In cases where there is more than one portal to the nest, the ventilatory activity will occur at each of these sites. Apparently every hornet worker engages in such ventilation at one time or another and their number at any one time is temperature-dependent, that is, the higher the ambient temperature the greater the number the hornets participating in ventilation. Indeed the correlation between the temperature and the ventilatory activity is positive and highly significant. [ABSTRACT FROM AUTHOR]

Two combined mechanisms on the hornet tarsus are adapted to attachment to the substrate: a friction-based (claws and spines) and an adhesion-based one (arolium). There are two ranges of substrate roughness optimal for attachment, either very smooth or very rough. There is an intermediate range of substrate grains of small but non-zero size, where both of these mechanisms fail. The optimal size of substrate grains for hornet grasping was 50–100 μm. Maximal hold to the substrate was achieved when surface irregularities were clamped between the claws of opposite legs. In such a position, the insect could withstand an external force which was almost 25 times larger than its own weight.The tarsal chain is an important part of the entire attachment mechanism. The articulations in the kinematic chain of tibia-tarsus-pretarsus are monocondylar. Three tarsal muscles and one head of the claw retractor muscle originate in the tibia. On pull to the retractor tendon, the tarsus bends in a plane. All elements of the tarsal kinematic chain have one active degree of freedom. The distance between the intertarsomeric articulation point and the tendon of the claw retractor (75–194 μm) corresponds to an efficiency of 1° per 1–3 μm of pulling distance travelled by the tendon. The claw turns about 1° per 4.3–5.0 μm of pulling distance travelled by the unguitractor.The arolium turns forward and downward simultaneously with flexion of the claws. The kinematic chain of the arolium lacks real condylar joints except the joint at the base of the manubrium. Other components are tied by flexible transmissions of the membranous cuticle.The walking hornet rests on distal tarsomeres of extended tarsi. If the retractor tendon inside the tarsus is fixed, passive extension of the tarsomeres might be replaced by claw flexion. Tarsal chain rigidity, measured with the force tester, increased when the retractor tendon was tightened. Probably, pull to the tendon compresses the tarsomeres, increasing friction within contacting areas of rippled surfaces surrounding condyles within articulations. [Copyright &y& Elsevier]

The relationship between reproductive status and body size in foundresses of Ropalidia plebeiana, an Australian endemic paper wasp forming huge aggregations of nests, was examined. Foundresses with developed ovaries (laying foundresses) in multifoundress colonies tended to be larger than foundresses in single-foundress colonies and foundresses with undeveloped ovaries (non-laying foundresses). However, the laying foundress was not always the largest of the foundresses in a colony. Body shape in foundresses varied according to simple allometric growth, while foundresses and first brood females diverged in their growth parameters in the preimaginal stage. [ABSTRACT FROM AUTHOR]

Abstract Mating frequency of Vespa analis queens and the genetic relatedness of their workers was analyzed by DNA microsatellite genotyping. Of 20 colonies studied, 18 had a queen inseminated by a single male and two had queens each inseminated by two males. The estimated effective number of matings was 1.05 ± 0.037 (mean ± SE), with 75–85% of the offspring of the two multiply mated queens sired by a single male. The pedigree relatedness between nestmate workers averaged over the 20 colonies was estimated to be 0.74 ± 0.008, almost identical to the predicted value of 0.75 for colonies headed by a singly mated queen. Multiple matrilines; that is, the presence of workers not related to the current queens, were detected in six colonies, suggesting that queen replacement occurred via usurpation of the founding queens in these six colonies. These results demonstrate that the kin structure of V. analis is similar to that reported in other vespid species. [ABSTRACT FROM AUTHOR]

Background: Venom sac extract of yellow jackets Vespula vulgaris was toxic in mice when injected intraperitoneally but not toxic when injected subcutaneously. Necropsy showed the toxicity to be an inflammatory response. Methods: Venom peptide and protein fractions were tested to identify the inflammatory components. The active components were tested to establish whether they might function as adjuvant for venom protein-specific antibody response. Results: Venom toxicity required the synergistic action of two venom components, a mast cell degranulating peptide mastoparan and phospholipase A1. Both components stimulated prostaglandin E[sub 2] release from murine peritoneal cells and macrophages. Mastoparan showed a weak activity to enhance IgE and IgG1 responses to a yellow jacket venom protein Ves v 5 in BALB/c mice. It was not possible to assess the adjuvant activity of phospholipase A1 because of its suppression of Ves v 5-specific response. Melittin, a mast cell degranulating peptide from bee venom, was inactive as an adjuvant for Ves v 5-specific response. Conclusion: Yellow jacket venom contains two inflammatory components, mastoparan and phospholipase A1. Our findings suggest that mastoparan can function as a weak adjuvant for TH2 cell-associated antibody response.Copyright © 2003 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

Abstract Morphological differences between queens and workers in an Australian swarm-founding paper wasp, Ropalidia romandi , were examined, providing detailed information on the nature of morphological caste differences in swarm-founding Ropalidia for the first time. Queens and workers differed in overall size and had divergent shapes, but differences were not always significant. In colonies in which queen–worker dimorphism was distinct, while the head width of queens was nearly the same as that of workers, queens were significantly larger in measures on the metasoma than workers. Plotting of metasomal variables against the mesosomal length and analysis of covariance showed that queen–worker morphological differentiation in R. romandi cannot be explained by simple linear allometric growth, suggesting that the two castes diverge in their growth parameters in the preadult stage. The pattern of morphological caste difference varied among colonies or was colony specific; there was no clear correlation between caste differences and colony cycle progression. These results suggest that morphological caste differences in R. romandi are determined genetically or by interactions between genetic background and intrinsic factors and/or external conditions. [ABSTRACT FROM AUTHOR]

Who reproduces in colonies of social insects is determined by some combination of direct competition and more peaceful convention. We studied these two alternatives in foundresses of the paper wasp, Polistes carolina, by examining two different contexts: what determines who becomes the dominant reproductive and what determines the amount of reproduction obtained by subordinates. The dominant queen on most nests was the foundress to arrive first, rather than the largest foundress, expected to be best at fighting. This suggests that dominance is initially determined by convention, although the persistence of some aggressive conflict throughout the foundress period suggests that this convention is not absolute. Attempts to explain the division of reproduction using several skew theories were generally unsuccessful. Skew was not correlated with relatedness, size differences, colony productivity, and challenges by the subordinate. P. carolina showed high constraints against solitary nesting, with a minority of females attempting to nest alone, and none succeeding. In this situation, most skew theories predict that group stability will be independent of relatedness, yet nearly all collected subordinates were full sisters to the queen. Reproductive partitioning in early P. carolina colonies may have more to do with enhancing worker production than with conflict over direct fitness. [ABSTRACT FROM AUTHOR]