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.     

Colony productivity and foundress behaviour of a native wasp versus an invasive social wasp

Abstract.  1. Colony productivity, prey utilisation, and foundress behaviour of a North American native wasp ( Polistes fuscatus ) versus an European invasive wasp ( Polistes dominulus ) were investigated in a controlled field experiment with optimal versus natural foraging conditions. Colonies with the optimal prey foraging conditions were provided with prey ad libitum within an enclosed area. The other colonies foraged in the adjacent field–woodland but had the same nest conditions as the other treatment.
2. When given prey ad libitum , both wasp species captured similar amounts of prey and the conversion to total offspring biomass was similar. But P. dominulus colonies produced 2.5 times the number of workers as P. fuscatus colonies, reflecting the smaller size of P. dominulus wasps.
3. Foundresses of P. dominulus were observed more often building or repairing the nest, thereby contributing to the production of colonies with twice as many cells as colonies of P. fuscatus . Foundresses of P. dominulus showed more acts of aggression toward workers than did P. fuscatus foundresses, which was not a function of adult density on the nest.
4. At the end of the experiment, P. dominulus colonies with optimal prey foraging conditions still had a high level of egg-laying and peaked in the number of pupae then, whereas egg-laying and the number of pupae per colony of the other treatments began to decline 2–3 weeks earlier. These results indicate that P. dominulus is more opportunistic than P. fuscatus , which may account in part for P. dominulus 's success as an introduced species in North America.     

Venom volatiles of the paper wasp social parasite <Emphasis Type="Italic">Polistes sulcifer</Emphasis> elicit intra-colonial aggression on the nest of the host species <Emphasis Type="Italic">Polistes dominulus</Emphasis>

Insect social parasites rely on host workers to rear and protect their own brood. To conquer a host colony, a parasite must overcome the defensive mechanisms of the host, often by exploiting its chemical communication system. A widespread strategy involves the production of specific allomones (the so-called “propaganda pheromones”) to facilitate the usurpation process by manipulating the defensive behavior of the host. Polistes sulcifer is the obligate and permanent social parasite of the congeneric paper wasp Polistes dominulus. In this study, we investigated if the venom volatiles, well known to be alarm pheromones in the host species, could be used by the parasite to manipulate the host defense. We thus performed laboratory bioassays, to evaluate the possible effect of the venom volatile compounds of the parasite on the host. Our results show that host colony members reacted to the venom volatiles extract of the parasite with an increase in intra-colonial aggression compared to the reaction induced by the venom volatiles extract of the host foundress. Besides, a re-analysis of previously published chemical data showed that the parasite venom volatiles profile differs from that of the host: the spiroacetals are absent, whilst the amides are very abundant in the parasite venom when compared with that of the host. Similar to other insect social parasites, Polistes wasp parasites might be able to increase their invasion success by using venom volatile pheromones to distract the host defenders.     

Alarm communication in <Emphasis Type="Italic">Ropalidia</Emphasis> social wasps

Summary Alarm pheromones, chemical substances produced by social insects to alert the colony to threat, are the principal means by which colony defence is co-ordinated. We present the results of a study on alarm behaviour in 5 swarming species of wasps belonging to the genus Ropalidia. These species show a remarkably efficient strategy of alarm communication, including visual display and attack synchronization. We show that pheromones released from the venom gland play an important role in alarm recruitment in species belonging to the Ropalidia flavopicta group, but not in Ropalidia sumatrae. We analysed the contents of the venom reservoirs content of four of the studied species by gas chromatography-mass spectrometry. Glands were found to contain a complex mixture of volatile compounds as well as spiroacetals of higher molecular weight. Interestingly, despite all species producing similar chemical compounds from the venom gland, these were found to elicit alarm behaviour in only those species that build nest envelopes, suggesting a link between chemical release of alarm behaviour and the evolution of nest architecture in Ropalidia wasps.Received 19 August 2003; revised 29 February 2004; accepted 10 March 2004     

Can venom volatiles be a taxonomic tool for Polistes wasps (Hymenoptera,Vespidae)?

In the present study, we investigated whether venom volatiles have a species-specific composition and could possibly be used to discriminate between related paper wasp species. We compared venom volatile chemical profiles, obtained through gas chromatography-mass spectrometry analyses, of three non-parasitic European Polistes species ( P. dominulus , P. gallicus and P. nimphus ). The results show that the venom volatile composition is indeed species specific and can thus be a useful systematic tool.     

Effects of human presence on two social wasp species

Abstract.  1. The effects of human presence on the behavioural response and productivity of a native wasp species of North America ( Polistes fuscatus ) and an invasive wasp species ( Polistes dominulus ) were examined in a field experiment. Over a 3-month period, colonies of each species were exposed to human presence at either low (biweekly) or high (daily) levels. At 2-week intervals, wasp response to human presence was recorded for all colonies via an index, ranging from no response (0) to an attack (4).
2. The levels of human presence had no effect on wasp response or colony productivity but foundresses of both species increased their response level the closer a human was to the nest. In addition, foundresses of P. fuscatus became more aggressive over time, corresponding to after adult offspring emerged, whereas foundresses of P. dominulus did not change their response level after the emergence of offspring.
3. The adult offspring of P. dominulus responded to human presence at a level similar to that of the foundress, whereas adult offspring of P. fuscatus exhibited a lower response to human presence than that of the foundress. The more distinctive aposematic coloration of P. dominulus may play a role in the behavioural differences between species; it may be more effective in deterring predators, and with the result that leaving the nest to defend it is less often necessary. Accordingly, the less aggressive response to human presence by P. dominulus may contribute to its success as an introduced species in North America.     

Rank integration in dominance hierarchies of host colonies by the paper wasp social parasite Polistes sulcifer (Hymenoptera, Vespidae)

In multiple-foundress nests of the wasp Polistes dominulus, dominance hierarchies are established among foundresses, and only the dominant (=alpha) individual lays eggs. The alpha female can be distinguished from subordinate females and workers on the basis of the proportions of some hydrocarbons present on the cuticle, suggesting that chemical signaling of her reproductive status could occur. P. dominulus is also the host species of the obligate social parasite Polistes sulcifer. After aggressively usurping host colonies and behaviorally replacing the host alpha female, parasites are characterized by a change in the proportions of their cuticular hydrocarbons to match that of the host cuticular profile at both species and colony levels. In the current study, we demonstrate that P. sulcifer queens also modify their cuticular hydrocarbon proportions after usurpation to match that of the host alpha female. Parasite females, therefore, acquire the dominant rank in host colonies both reproductively and chemically by mimicking the typical alpha profile of the host. Parasite females were not able to fully inhibit ovary development in host foundresses, and 10 days after usurpation, parasites, alpha and beta foundresses show similar chemical profiles and ovarian development.     

Dynamics of cuticular chemical profiles of Polistes dominulus workers in orphaned nests (Hymenoptera, Vespidae)

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.     

The chemical strategies used by Polistes nimphus social wasp usurpers (Hymenoptera Vespidae)

Polistes foundresses can behave as facultative social parasites when, instead of founding their own nest, they usurp colonies of the same or a different species and temporary use the host workforce to raise their own brood. Conspecific usurpation appears to be common among Polistes wasps, but nothing is known about the mechanisms that these facultative social parasites use to have themselves accepted within usurped colonies. Using behavioural tests, we studied the chemical strategies employed by females of Polistes nimphus when they behave as facultative social parasites in colonies of the same or of a different species. We hypothesized that usurpers would mark host nests with their own odours and/or acquire host nest odours in order to camouflage their real identity from host workers. Our results indicated that P. nimphus usurpers used different chemical strategies depending on host nest species: they acquired conspecific host odours but marked heterospecific host combs with their own odours.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 505–512.     

Recognition of social parasites as nest-mates: adoption of colony-specific host cuticular odours by the paper wasp parasite Polistes sulcifer.

Colonies of the polistine wasp Polistes dominulus are parasitized by the permanent worker-less social parasite Polistes sulcifer. After usurpation of the host colony, parasite females are characterized by a change in the relative proportions of their cuticular hydrocarbons to match those of the host species. In this paper we present evidence from field data and laboratory experiments that P. sulcifer females adopt a colony-specific host odour that facilitates their acceptance by host females of the usurped colony. Presentation experiments demonstrate that parasite females are recognized as foreign individuals by workers of other parasitized nests. We show that the modification of parasite cuticular compounds is sufficient for this recognition. This provides evidence that, after invasion, P. sulcifer queens do not require appeasement or propaganda substances for their acceptance by host colonies. Furthermore, multivariate discriminant analysis of the cuticular hydrocarbon proportions of the parasites after usurpation assigns the parasites together with P. dominulus females of their own host colony. To the authors' knowledge, this is the first confirmation that social parasites adopt colony-specific host odours.     

Ectal mandibular gland in Polistes dominulus (Christ) (Hymenoptera, vespidae): ultrastructural modifications over the secretory cycle

An ultrastructural study was carried out on the secretory activity of the ectal mandibular gland in the wasp Polistes dominulus (foundress and worker females as well as males). Secretory activity in foundresses proceeds slowly during hibernation and early spring, becoming prominent in late spring and then falling sharply during the summer. This sequential pattern of ultrastructural modifications follows a functional, annual cycle. However, by comparing the subcellular changes in the gland with colonial development, it appears that secretory activity fits in with the specie's social cycle rather than merely following the seasons. The highest levels of secretory activity correspond to the early, critical breeding phases, while activity slows down with an increase in colony protection, based on both primary (passive) and secondary (active) defenses, with the emergence of the workers. These correlations suggest that the ectal mandibular gland secretory product in P. dominulus is involved in chemical nest defense.     

The morphology of Van der Vecth's organ as a tool to measure caste dimorphism in Polistes paper wasps: a comparative approach

Given the centrality of chemical communication in social insects, there are many selective pressures acting on morpho‐functional traits that mediate chemical pheromones. On the last gastral sternite of Polistes females, there is an important exocrine surface secreting chemical pheromone, named Van der Vecht's organ. It is involved in chemical defence of the nest, in rank and nestmate recognition, preventing workers from direct reproduction. Allometric differential growth of phenotypic traits between castes of social insects is generally considered as an indication of incipient physical castes. European Polistes present different nesting strategies and reproductive choices. Here, we carry out a comparison of Van der Vecht's organ size between castes of four European Polistes to provide a general measure of dimorphism. We show that Van der Vecht's organ of Polistes dominula and Polistes nimphus foundresses shows an allometric development being enlarged with respect to workers. Otherwise, no allometries have been highlighted in the other two studied species (i.e. Polistes associus and Polistes biglumis). Therefore, our data show that neither rigid monogyny nor specific nesting habits foster the evolution of true morphological castes in primitively eusocial taxa. Thus, at least two other species of Ezuropean Polistes show real evidence of incipient morphological castes.     

Isolation, cloning and characterization of Polistes dominulus venom phospholipase A1 and its isoforms

Polistes dominulus is known to be an allergenically important social wasp. Its venom has four major allergens (ziv. phospholipase A1, hyaluronidase, antigen 5 and a serine-protease). Amino acid sequences of its serine-protease and Antigen 5 have been published. In this paper, the partial amino acid sequence of its venom phospholipase native protein is reported. Also, we give an account to the complete nucleotide sequence of Polistes dominulus venom phospholipase A1 gene, its isoforms and their complete deduced amino acid sequences. Their similarity to the other phospholipases A1 of the family: Vespidae is discussed.     

Not only cuticular lipids: first evidence of differences between foundresses and their daughters in polar substances in the paper wasp Polistes dominulus

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 2679 Da, 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.     

Workers of a polistes paper wasp detect the presence of their queen by chemical cues

Differences in long-chain hydrocarbon mixtures among reproductive and nonreproductive individuals have been often revealed in social insects. However, very few papers demonstrated that these signatures actually act as contact pheromones used by nonreproductive to recognize the presence of a related queen in the colony. Cuticular and glandular hydrocarbons of Polistes paper wasps have been extensively studied, but, until now, the perception and recognition of such cues was not demonstrated. In this paper, we show, for the first time in Vespidae, that Polistes gallicus workers distinguish nestmates from alien individuals and queens from workers by the hydrocarbon mixtures of the Van der Vecht organ secretion (VVS). We also demonstrated that stroking behavior (a peculiar behavior of Polistes by which queens probably lay VVS on the nest) acts as an inhibitor of ovarian development in workers.     

Cuticular hydrocarbons rather than peptides are responsible for nestmate recognition in Polistes dominulus

A colony of social insects is like a fortress where access is allowed only to colony members. The epicuticular mixture of hydrocarbons has been widely reported to be involved in nestmate recognition in insects. However, recent studies have shown that polar compounds (mainly peptides) are also present, mixed with hydrocarbons, on the cuticle of various insects, including the paper wasps of the genus Polistes. As these polar compounds are variable among Polistes species and are perceived by the wasps, this cuticular fraction could also be involved in nestmate recognition. Through MALDI-TOF (Matrix-Assisted Laser Desorption Ionization Time of Flight) mass spectrometry analysis, we assessed, for the first time, the intercolonial variability of the cuticular polar fraction of Polistes dominulus in order to evaluate its reliability as source of nestmate recognition cues. We then tested through behavioral assays the importance of the 2 isolated fractions (apolar and polar) in nestmate recognition by presenting them separately to colonies of P. dominulus. Our results showed that the cuticular polar compounds are not colony specific and they are not used by paper wasps to discriminate nestmates from non-colony members. On the contrary, we confirmed that the isolated cuticular hydrocarbons are the chemical mediators prompting nestmate recognition in paper wasps.     

Variation in cuticular hydrocarbon signatures, hormonal correlates and establishment of reproductive dominance in a polistine wasp

In many social insects the relationship between reproductive dominance and physiological correlates is poorly understood. Recent evidence now strongly suggests that cuticular hydrocarbons are important in reproductive differentiation in these societies where they are used as signals of ovarian activity in reproductive females. In this study we investigated the relationship between reproductive dominance, size of the corpora allata (CA, producer of Juvenile Hormone, JH) and the proportions of cuticular hydrocarbons present on the cuticle in overwintering foundresses and both associative (polygynous) and solitary (monogynous) pre-emergence colonies of the social wasp Polistes dominulus. Size of the CA was positively correlated with ovarian development in polygynous colonies. In contrast, solitary foundresses possessed significantly smaller CAs than dominant foundresses from polygynous nests, yet ovarian activity was similar for both female types. CA size variation was associated with variation in cuticular hydrocarbon proportions. Overwintering, solitary, dominant and subordinate (from associative nests) females all possessed distinctive cuticular chemical profiles revealed by multivariate discriminant analyses. Our data indicate that the social environment strongly affects reproductive physiology in this wasp, and we discuss the role of cuticular hydrocarbons in reproductive signaling in P. dominulus and other social insects.     

Nourishment affects colony demographics in the paper wasp Polistes metricus

Abstract  1. Colony survivorship and numbers of nest cells, pupae, and adult females were monitored throughout the nesting season for a cohort of 78 colonies of the paper wasp Polistes metricus Say. Thirty-nine colonies received a twice-weekly nourishment supplement of honey during pre-emergence and early emergence phases of the colony cycle; 39 colonies were unsupplemented controls.
2. Colony survivorship was unaffected by the supplemental nourishment. Loss of colonies to predation differed among three sites but was unaffected by supplementation.
3. Honey-supplemented colonies constructed more nest cells than did control colonies but this effect was not expressed until after supplementation had ceased.
4. Honey-supplemented colonies produced more pupae than did control colonies but the number of adult females at nests did not differ between supplemented and control colonies. Because honey-supplemented colonies had more offspring but fewer of them remained as workers at the nest, honey supplementation led to a lower frequency of workers and corresponding higher frequency of reproductives than in control colonies.
5. In a second year of study, colony survivorship and numbers of nest cells, pupae, and adult females were monitored from late pre-emergence until the end of the nesting season for a cohort of 32 colonies of Polistes metricus . In 16 colonies, trophallactic saliva was taken from final-instar larvae on nine dates in the late pre-emergence and early emergence periods; 16 colonies served as controls.
6. Saliva-diminished colonies had lower survivorship, fewer nest cells, fewer pupae, and fewer adult females at the nest than did control colonies.
7. These results show that variation in nourishment in the early to mid phases of the colony cycle can have significant effects on the subsequent colony demographics of Polistes metricus paper wasps.     

Orientation and Communication in the Neotropical Ant Odontomachus bauri Emery (Hymenoptera,Formicidae, Ponerinae)

The Neotropical species Odontomachus bauri employs canopy orientation during foraging and homing. An artificial canopy pattern above the ants is much more effective as an orientation cue than horizontal landmarks or chemical marks. However, both horizontal visual cues and chemical marks on the ground can serve in localizing the nest entrance. Successful O. bauri foragers recruit nestmates to leave the nest and search for food. However, the recruitment signals do not contain directional information. Antennation bouts and pheromones from the pygidial gland most likely serve as stimulating recruitment signals. Secretions from the mandibular and poison gland elicit alarm and attack behavior.     

Cryptic castes,social context and colony defence in a social bee,Tetragonula carbonaria

Division of labour in social insect colonies is facilitated in two ways: through temporal sharing of tasks or by morphologically specialised castes. In casteless species, colony defence is maintained by morphologically indistinct workers, who lack the obvious defensive specialisation of polymorphic species. Discrimination of intruders is carried out via antenna, which also detects defensive social cues such as alarm pheromones. Despite their functional importance however, antennal morphology is rarely considered in studies of nestmate recognition. We investigated antennal morphology and the necessity of social cues in mediating defensive behaviour across differentially tasked workers of a casteless social bee, Tetragonula carbonaria. Our results suggest that the current understanding of division of labour in casteless worker species remains poorly understood, with differences in antennal morphology and aggression creating morphologically and behaviourally distinct ‘cryptic castes’. Further, we found that defensive behaviour was only elicited near nest odours, highlighting the importance of mediating aggression among workers.