Colony founding in the primitively eusocial wasp, Ropalidia marginata (Hymenoptera: Vespidae)

Abstract.

• 1 In a 16-month study in Bangalore, India, about 35% of the newly founded colonies of Ropalidia marginata were single foundress colonies and the remainder were multiple foundress colonies with two to twenty-two individuals.
• 2 Larger colonies did not have a significantly higher per capita productivity, did not produce significantly heavier progeny and did not produce them significantly faster than smaller colonies did.
• 3 Predation by the hornet Vespa tropica appeared to be independent of group size.
• 4 Single foundress colonies failed more often but not often enough to make them have a lower average per capita productivity, compared to multiple foundress colonies.
• 5 Some of the advantages of multiple foundress associations came from the greater predictability of their attaining the mean per capita productivity, the relatively lower rates of usurpation experienced by them compared to single foundress colonies, and the opportunities provided by queen turnovers for workers to become replacement queens and gain direct individual fitness.

     

Mating structure and genetic relatedness among gynes in the primitively eusocial wasp Polistes snelleni (Hymenoptera: Vespidae)

Mating structure and genetic relatedness among gynes (potential reproductive females) of the paper wasp Polistes snelleni were 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.     

Absence of chemical alarm in a primitively eusocial wasp (Belonogaster petiolata,Hymenoptera: Vespidae)

Summary The hypothesis thatBelonogaster petiolata (fam. Vespidae) is able to communicate alarm chemically, using odours released with the venom, was tested in bioassays involving presentation of artificial targets to a wasp colony, simultaneously with crushed venom apparatuses. The odour of venom did not lower the threshold of attack and visual stimuli alone (particularly a black, moving object) were sufficient to release attack. Venom odour on a previously stung target probably does not play a role in focusing further attacks on such a target. The results therefore support the null hypothesis that a venom-based alarm pheromone is absent in this species.     

Behavioral role differentiation in the primitively eusocial waspBelonogaster juncea juncea (Hymenoptera: Vespidae)

With a view toward describing behavioral variability among individuals of the primitively eusocial speciesBelonogaster juncea juncea, we recorded the time-activity budget spent on five behavioral categories (foraging, building, feeding, inactivity, and reproduction) by 52 individuals belonging to four postemergence colonies. A principal-components analysis coupled with a hierarchical cluster analysis enabled us to discern four behavioral roles. The reproductive role is reserved to one individual (functional monogyny) and the workers’ role is differentiated into foragers, builders, and guards. The foragers are females that spend the most time (82.6% of their time) foraging, supplying the nest with prey load and liquid matter. The builders are individuals (with 41.5% of their time off the nest) that tend to bring pulp into the nest and then undertake building activities. The guards are those females that spend the most time (79.7% of their time) being inactive on the nest.     

Reproductive and worker castes in the primitively eusocial wasp Belonogaster petiolata (DeGeer) (Hymenoptera: Vespidae): evidence for pre-imaginal differentiation

Until recently, morphological differences between castes in independent-founding polistine wasps were considered absent. This paper investigates the extent of morphological and physiological differences between reproductive (foundress and gyne) and worker castes of Belonogaster petiolata, and tests the hypothesis that caste differentiation in this species occurs pre-imaginally.Foundresses were significantly larger than workers, to the extent that foundress/worker ratios were comparable with those between queens and workers in some swarm-founding Polistinae. Early emerging workers were small, but body size increased over the colony cycle such that late-season workers were similar in size to gynes. In proportion to body size, workers possessed broader heads while foundresses and gynes had broader thoraces and gasters. All queens, 98% of subordinate foundresses, and 95% of over-wintering gynes were inseminated. Workers were never inseminated and lacked mature ovaries in colonies with active queens. However, in the absence of the queen (and other foundresses), 11% of workers developed mature ovaries. Ovarian size and fat content of foundresses and gynes was significantly greater than that of workers. The differences in external morphology and reproductive physiology between castes support the hypothesis that differentiation occurs pre-imaginally. However, imaginal factors, in particular social dominance of the queen, maintain the reproductive subordinance of workers.     

Dominance behaviour and regulation of foraging in the primitively eusocial wasp Ropalidia marginata (Lep.) (Hymenoptera: Vespidae)

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.     

Factors affecting the acceptance of alien conspecifics on nests of the primitively eusocial wasp,Ropalidia marginata (Hymenoptera: Vespidae)

In the primitively eusocial wasp, Ropalidia marginata,individual females are known to drift from one newly founded nest to another. In the laboratory, young (<6- to 8-day-old) alien wasps are accepted onto unrelated colonies, while older (>6- to 8-day-old) wasps are not. Here we have investigated the factors that could influence the acceptance of foreign conspecifics onto unrelated nests. Individually marked wasps of different ages, isolated immediately after eclosion from the natal nest and from each other, were introduced onto unrelated recipient nests. Considered separately, both age and ovarian condition seemed to influence the probability of acceptance as well as the levels of aggression and tolerance received by the introduced wasps. However, partial correlation analysis and multiple regression analysis indicated that only age had a direct influence and that the ovarian condition acts only through age, a variable with which it is highly correlated. The observed acceptance of young aliens and rejection of old aliens are less likely to be due to the perception of older wasps as a reproductive threat rather than some age-related factor, other than ovarian condition, for example, the relative ease with which younger wasps can be molded into desired roles.     

The role of larval nutrition in pre-imaginal biasing of caste in the primitively eusocial wasp Ropalidia marginata (Hymenoptera: Vespidae)

Abstract.

• 1 When freshly eclosed females of the primitively eusocial wasp Ropalidia marginata (Lep.) are isolated, only about 50% of them build nests and lay eggs thereby suggesting a pre-imaginal biasing of caste.
• 2 Wasps that lay eggs take a very variable amount of time after eclosion to start doing so.
• 3 Females eclosing from nests where larvae are fed at a relatively higher rate are more likely to become egg-layers and are likely to take less time after eclosion to begin to lay eggs.
• 4 Thus, both forms of pre-imaginal biasing of caste, namely, differences in egg laying capacity and differences in the time taken to attain reproductive maturity, appear to be influenced by larval nutrition.

     

Unmated queens in the primitively eusocial waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Summary In the primitively eusocial tropical waspRopalidia marginata, five out of eleven colonies studied had an unmated female as their queen. In two colonies this was the case despite the presence of another mated individual in the colony. We found no detectable differences between colonies with unmated queens and those with mated queens. We argue that in species such asR. marginata, where intracolony relatedness is expected to be low and where sociality is likely to be maintained because several individuals have opportunities for direct reproduction in the future, individual selection is likely to override the good of the colony and lead to such phenomena as that of unmated queens.     

Size-dependent reproductive dominance in foundresses of Ropalidia plebeiana, an Australian paper wasp forming nest aggregations (Hymenoptera: Vespidae)

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.     

Queen succession in the primitively eusocial tropical waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Females of the primitively eusocial wasp Ropalidia marginata can be classified into three behavioral groups: Sitters, Fighters, and Foragers. It has been speculated that both Sitters and Fighters may be hopeful queens and that the Foragers may have little or no opportunities for direct reproduction. Here we show that in 9 of 12 queen-removal experiments where such a behavioral differentiation could be discerned, the individual that became a queen (the potential queen) was a Sitter in 6 cases, a Fighter in 2 cases, and a Forager in only 1 case. Although potential queens spent significantly more time absent from the nest and showed significantly higher rates of dominance behavior compared to the mean values for nonqueens in their colonies, they were intermediate with respect to all behaviors and age when compared to the range of values for nonqueens in their colonies. Potential queens were not necessarily the highest-ranking individuals among the nonqueens. The pattern of queen succession in this species appears to be quite different from the temperate pattern, where an old and active forager of high dominance rank is the potential queen. Although somewhat similar to the tropical pattern of a relatively younger female that has performed relatively little foraging being the potential queen, it is perhaps more accurate to describe the potential queens of R. marginata as unspecialized intermediates.     

Dual mimicry in the dimorphic eusocial wasp Mischocyttarus mastigophorus Richards (Hymenoptera: Vespidae)

The eusocial vespid wasp Mischocyttarus mastigophorus exhibits two colour morphs, with males and females of each morph co-occurring at Monteverde, Costa Rica. Each morph closely resembles a different sympatric species of swarm-founding wasp in the genus Agelaia. We propose that the Agelaia species are models for a dual mimicry system. The Agelaia species (A. yepocapa , mimicked by the M. mastigophorus pale morph, and A. xanthopus , mimicked by the M. mastigophorus dark morph) are locally abundant wasps with large, aggressively defended colonies. The mimic and models are restricted to high-elevation habitat in the Monteverde area, and the elevational ranges of both Agelaia species partially overlap the elcvational range of M. mastigophorus. Relative frequencies of the M. mastigophorus colour morphs vary with elevation, with the pale morph predominating at lower elevations. Elevational differences in the relative abundances of the Agelaia species suggest that the models act as a selective force maintaining the M. mastigophorus colour polymorphism at Monteverde. Mischocyttarus mastigophorus overlaps only A. xanthopus in the northern part of its range (S. Mexico), and overlaps only A. yepocapa in the southern part of its range (Ecuador). We hypothesize that the M. mastigophorus colour morphs evolved in allopatry and later came into contact in Central America. Appropriate high-elevation habitat for cloud forest species is distributed as discrete patches in Central America and Northern South America. The island-like nature of suitable habitat may favour the isolation and rapid evolutionary diversification of vespid species that are restricted to high elevations in the Neotropics.     

Evidence for multiple mating in the primitively eusocial waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Asymmetries in genetic relatedness created by haplodiploidy have been considered to be crucially important for the evolution of worker behaviour in Hymenoptera. Multiple mating by the queens destroys this asymmetry and should make kin selection less powerful. The number of males that social insect queens mate with is thus of considerable theoretical interest especially in primitively eusocial species. The results presented here provide evidence for multiple mating by foundresses of the primitively eusocial waspRopalidia marginata (Lep.)     

Division of labor in post-emergence colonies of the primitively eusocial waspPolistes instabilis de Saussure (Hymenoptera: Vespidae)

Polyethism was quantified in post-emergence colonies of the primitively eusocial wasp,Polistes instabilis, and compared to polyethism in a sympatric advanced eusocial wasp,Polybia occidentalis. Like P.occidentalis, P. instabilis foragers collected food (nectar and prey) and nest materials (wood pulp and water).P. instabilis foragers showed some evidence of specialization with respect to which materials they gathered, but most foragers, divided their effort among food and nest materials, a pattern that is rarely seen inP. occidentalis. In colonies of both species, more foragers collected nectar than any other material; in contrast, most water foraging was performed by one or two workers. Upon returning to the nest,P. instabilis foragers gave up part or all of most nectar, prey, and pulp loads to nestmates, while water was rarely partitioned. Prey loads were most likely to be given up entirely.P. instabilis workers show evidence of conflict over the handling of materials at the nest. The frequency with which workers took portions of nectar loads from forgers was positively correlated with their frequency of aggressive dominant behavior, and with their frequency of taking other foraged materials. Compared to polyethism inP. occidentalis P. instabilis showed less individual specialization on foraging tasks and less partitioning of foraged materials with nestmates, suggesting that these characteristics of polyethism have been modified during the evolution of advanced insect societies.     

Worker biting interactions and task performance in a swarm-founding eusocial wasp (Polybia occidentalis, Hymenoptera: Vespidae)

Workers in many insect societies interact via body contact withtheir nest mates, and social biting and other forms of contactmay play a general role in regulating task performance. HereI present evidence that social biting affects task performancewithout direct reproductive conflict in Polybia occidentalis,a swarm-founding eusocial wasp. Polybia occidentalis workersengaged in social biting with nest mates. Most workers thatwere active on the nest surface participated in biting interactions,but individuals differed significantly in their rates of biting and of being bitten. Rates of being bitten corresponded withnonreproductive task performance: more biting was directedat foragers than nonforagers, and foraging rates were correlatedwith rates of being bitten. Furthermore, some on-nest workersinitiated foraging activity immediately after they were bitten.Together these patterns suggest that social biting influencesforaging rates by increasing workers' probabilities of leavingthe nest. Variation in biting rates did not correspond withdifferences in reproductive physiology: highly active bitersand recipients did not differ in body size or in ovary development.In P. occidentalis and in other eusocial insects with largeworker forces, biting and other types of social contact amongworkers may regulate task performance independently of directreproductive competition.     

Demography of the Okinawan eusocial wasp Ropalidia fasciata (Hymenoptera: Vespidae) II. Effects of foundress group size on survival rates of colonies and foundresses, and production of progeny

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.