Central place water collection in a Japanese paper wasp, Polistes Chinensis antennalis

The water collection behaviour of colony foundresses of a Japanese paper wasp, Polistes chinensis antennalis, was recorded. A single round-trip of water collection consists of (1) an outward flight, (2) water-sucking, and (3) a homeward flight. The rate of water intake decreased as the length of water-sucking increased. There was a positive correlation between the duration of the outward flight and that of water-sucking. The variance of time for outward flights was larger than that for homeward flights. The data were consistent with the predictions of the central place foraging model of Orians & Pearson (1979).     

On “intercolonial” cannibalism in Japanese paper wasps,Polistes chinensis antennalis Pérez andP. Jadwigae dalla torre (hymenoptera: Vespidae)

Summary Foundresses of two species of Japanese paper wasps,Polistes chinensis antennalis andP. jadwigae, attacked other colonies of the same species. A foundress ofP. chinensis antennalis visited two nests of the same species, and ate larvae from them, while two foundresses ofP. jadwigae each visited a nest of the same species, eating larvae and pupae even when the foundress of the attacked nest was on her nest. In addition, a foundress ofP. jadwigae distributed flesh balls thus obtained among their larvae. Discussion was made on the adaptive significance of the inter-colonial cannibalism. It was considered that, at first, it increases the foraging efficiency and secondly it plays a role in regulating population density.     

Construction of Nest Defensive Structure According to Offspring Value and Its Effect on Predator's Attack Decision in Paper Wasps

Various organisms are known to build nests with defensive structures to protect their offspring from predation, but our understanding of plasticity in the nest structure remains poor. In this study, we investigated whether a paper wasp, Polistes chinensis antennalis, adjusted the construction of nest defensive structure according to the value of their offspring, and we also analysed the effect of adjusting the construction of the structure on predator's decision to attack. P. chinensis antennalis foundresses start a colony and maintain her nest alone until the emergence of workers. During this stage, foundresses often construct a defensive structure on cocoon caps of pupae using nest materials (pulp), which prevents predation of pupae by conspecifics from other nests. The value of pupae to the foundress varies among those in a nest, where the value is higher in pupae that spun the cocoon (and initiated pupation) earlier than other pupae in the nest. From field observations, we found that foundresses constructed a larger pulp structure on the cocoons of pupae that cocooned earlier in the order of cocoon spinning, even after considering confounding factors. We also found that the probability of a pupa being attacked by conspecific intruders decreased with the size of pulp structure on the cocoon. This indicates that intruders avoid attacking cocoons with larger pulp structures. Our study indicates that foundresses adjust the construction of nest defensive structures according to their offspring value, and this allows them to protect the high‐value offspring efficiently and effectively.     

Behavioral ecology of Japanese paper wasps,Polistes spp. IV. Comparison of ethograms between queens and workers ofP. chinensis antennalis in the ergonomic stage

Behavioral catalogues, transition matrices between 2 consecutive behavioral acts, and the frequency distribution of time spent in each behavioral act were described for queens and workers of a Japanese paper wasp,Polistes chinensis antennalis in the ergonomic stage (the period from the emergence of the first worker to the emergenece of the first reproductive). The results were; (1) The whole behavior repertoire of queens was completely included in that of workers. Workers oviposited frequently in queen-right colonies. (2) The proportion of off-nest activities to the number of total acts for workers was higher than that for queens. The proportion of time spent off the nest in workers was 5 times as high as that in queens. (3) There was no significant difference in the proportion of number of oviposition to the total number of acts between queens and workers. The oviposition rate was 0.06 eggs/h for workers and 0.37 eggs/h for queens. Queens made more dominance behavior and workers more subordinated behavior. Queens made abdomen wagging more frequently than workers. (4) For behavioral acts common in both queens and workers, the mean duration of a specific act was approximately equal for both. The mean duration of unsuccessful flesh collection for workers was 3.4 times as longs as that for queens. Workers did not search for flesh in 10.1% of duration of flesh collections in the field. (5) The pattern of transitions between 2 consecutive behavioral acts in workers was similar to that in queens. (6) Alien workers approaching a nest with a load were more often accepted by the nest occupants than alien workers with no load. Alien workers with no load were not accepted as often as a legitimate nestmates with no load. The reproductive division of labor between queens and workers inP. chinesis antennalis was incomplete. This suggests worker-queen conflict in different reproductive strategies among colony members. Several behaviors were consistent with the view that they are the behavioral expression of this conflict.     

Effect of food availability on larval cannibalism by foundresses of the paper wasp <Emphasis Type="Italic">Polistes chinensis antennalis</Emphasis>

Cannibalism is a behavioral trait seen in a number of species and can be favored when the fitness gain outweighs the cost. In social wasps, field studies have suggested that food limitation causes larval cannibalism as a compensation of a meal for adult wasp and/or earlier production of the first workers, but experimental studies for the effect of food availability on larval cannibalism have been very scarce. Hence we examined whether Polistes chinensis antennalis foundresses exhibit a higher level of larval cannibalization under food-limited conditions than under food-available conditions. Larval cannibalization occurred frequently under both prey- and honey-limited conditions; however, young larvae were more frequently cannibalized than old larvae in the prey-limited colonies, while old larvae were more frequently cannibalized than young larvae in the honey-limited colonies. The number of days between colony initiation and the emergence of the first workers was not significantly different between the prey-limited and prey-available colonies, suggesting that under prey-limited conditions P. chinensis antennalis foundresses tend to cannibalize their young larvae and feed probably the flesh of these cannibalized larvae to old larvae in order to ensure the rapid production of the first workers. On the other hand, the number of days between colony initiation and the emergence of the first workers was longer in the honey-limited colonies than in the honey-available colonies. We propose explanations for the first time for why the foundresses in the honey-limited conditions cannibalized their old larvae more frequently than their young larvae in spite of this resulting in the delayed emergence of the first workers.     

Mothers vigilantly guard nests after partial brood loss: a cue of nest predation risk in a paper wasp

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Nesting biology of Asian paper wasps Polistes chinensis antennalis Pérez,and Australian paper wasps P. humilis (Fab.) (Hymenoptera: Vespidae) in northern New Zealand

Abstract

We recorded the numbers of cells, and where possible distinguished between cells containing pupating larvae and vacated cells, from 585 paper wasp nests from the northern North Island, New Zealand, plus nest site characteristics of 540 of these nests. Nests of Polistes chinensis antennalis and P. humilis developed at similar rates in early summer. P. c. antennalis nests were larger at the Post‐emergence stage than those of P. humilis, and contained more vacated cells but less capped cells. All of the P. c. antennalis nests had reached the Post‐emergence stage by February in Northland, but not in the other regions. P. c. antennalis nests in the Post‐emergence stage were larger in Northland than further south, and contained the most capped or vacated cells. Nests of both species were usually found in northern‐facing sites. Substrate did not affect nest size. Differences between the species in nest sites included greater use of manmade structures by P. c. antennalis; the use of leaves by P. humilis only; and a higher average nest site height in P. humilis. These differences in nest site selection may reduce competition between the species.     

Social behavior of early emerging males of a Japanese paper wasp,Polistes chinensis antennalis (Hymenoptera: Vespidae)

Summary Five cases of the early emergence of males in a Japanese paper wasp,Polistes chinensis antennalis, in which male emerged together with the first group of workers, were described. In one case of the five where the queen disappeared before the emergence of male, worker(s) produced female offspring. The frequency of the nests where the early emergence of male was observed was 16.7% (5/30 nests). In two colonies, worker(s) and/or queen chased off males. But in an orphan nest where worker(s) produced female offspring, the dominance order among workers which was similar to that of colony without male was observed. The significance of the early emergence of male in the social evolution of wasps was discussed.     

Why Wasp Foundresses Change Nests: Relatedness,Dominance, and Nest Quality

The costs and benefits of different social options are best understood when individuals can be followed as they make different choices, something that can be difficult in social insects. In this detailed study, we follow overwintered females of the social wasp Polistes carolina through different nesting strategies in a stratified habitat where nest site quality varies with proximity to a foraging area, and genetic relatedness among females is known. Females may initiate nests, join nests temporarily or permanently, or abandon nests. Females can become helpers or egglayers, effectively workers or queens. What they actually do can be predicted by a combination of ecological and relatedness factors. Advantages through increased lifetime success of individuals and nests drives foundresses of the social wasp Polistes from solitary to social nest founding. We studied reproductive options of spring foundresses of P. carolina by monitoring individually-marked wasps and assessing reproductive success of each foundress by using DNA microsatellites. We examined what behavioral decisions foundresses make after relaxing a strong ecological constraint, shortage of nesting sites. We also look at the reproductive consequences of different behaviors. As in other Polistes, the most successful strategy for a foundress was to initiate a nest as early as possible and then accept others as subordinates. A common feature for many P. carolina foundresses was, however, that they reassessed their reproductive options by actively monitoring other nests at the field site and sometimes moving permanently to new nests should that offer better (inclusive) fitness prospects compared to their original nests. A clear motivation for moving to new nests was high genetic relatedness; by the end of the foundress period all females were on nests with full sisters.     

Prey capture and caste-specific payload capacities in the European paper wasp Polistes dominulus

Here, we present a detailed ethogram for the foraging behavior of the eusocial paper wasp, Polistes dominulus. The animal’s foraging process can be sub-divided into four main stages: (1) approach, (2) attack, (3) butchering, and (4) balling. Although considerable behavioral variation exists within each stage, an analysis of more than 20 individual foraging bouts reveals a single “common path” leading from prey approach to its transportation back to the nest; elucidating this path and separating it into distinct stages is useful when identifying targets for future ethological studies. Of particular interest here is the balling phase, i.e., the preparation of a bolus of flesh for transportation back to the colony. Using an experimental approach, we show that foundresses can carry significantly heavier payloads than workers, suggesting a foraging advantage during the initial stages of colony development. We also show that wasp body mass is significantly positively correlated with payload capacity in foundresses, a relationship not seen among workers or late reproductives. This correlation suggests a beneficial adaptation of foundresses for combating early season pressures associated with the foundation of a new colony.     

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.     

Social behaviour of a subtropical paper wasp,Ropalidia fasciata (F.): Field observations during founding stage

Field studies showed that the percentage of foundation by multiple foundresses was remarkably high (34.5%) inRopalidia fasciata in Okinawa. Although one of the foundresses of the association usually remained on the nest and others tended to do extranidal activities, the dominance-subordination relation among cofoundresses seemed to be far milder than that reported inPolistes spp. of Europe and North America. Dominance relation sometimes reversed. All the foundresses could receive a share of food and feed larvae. It was suggested that a large risk of parasitism and predation should favour the trait to join foundress associations inR. fasciata.     

Nest site selection by fire ant queens

Summary We examined nest site selection by foundresses of the polygyne form ofSolenopsis invicta. In the laboratory, foundresses were allowed to nest in control soil vs. soil inoculated with refuse from a colony ofPheidole dentata, a natural enemy. In a second experiment foundresses chose between control andS. invicta soil. More foundresses (100%) avoided the soil of a natural enemy than the soil of a conspecific colony (79%) that may accept them into the nest. Foundresses from this polygyne ant appear to avoid other colonies in general, but avoid heterospecific colonies more.     

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

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

Early male production is not linked to a reproductive strategy in the Japanese paper wasp,<Emphasis Type="Italic"> Polistes chinensis antennalis</Emphasis> (Hymenoptera: Vespidae)

Diploid males in haplo-diploid insects are sterile, because they produce diploid sperm. Our previous report revealed that early males of the Japanese paper wasp Polistes chinensis antennalis are diploid but did not reveal how often this occurs. We analyzed the genotypes of early males using six microsatellite markers. Two of the 41 early males (5%) from six colonies were haploid, but the other males were diploid. This evidence suggests that we can ignore the reproductive success of the early males of P. chinensis antennalis.     

Nest Hydrocarbons as Cues for Philopatry in a Paper Wasp

Philopatric behavior has been demonstrated in a wide taxonomic spread of animals. In temperate environments, overwintered Polistes wasp foundresses often return to their natal nest prior to initiating colony construction. Previous research has shown that these spring foundresses can identify the natal nest in the absence of landmark and gross morphological cues. Hydrocarbons are essential recognition cues for Polistes nest and nestmate discrimination, but cuticular hydrocarbon profiles can become homogenized when foundresses overwinter in mixed colony groups. We examined the hydrocarbon profiles of Polistes dominulus foundresses and nests before and after an overwintering period, and found that the hydrocarbon profiles of nests remain unique over time and that this uniqueness is influenced by the original foundresses. Our data raise the possibility that in returning to the natal nest, foundresses reacquire their colony‐specific signature, which may play a role in the formation of cooperative associations.     

Foraging behavior of individual workers and foraging dynamics of colonies of three Sumatran stingless bees

The foraging behavior of three stingless bees, Trigona (Tetragonula) minangkabau, T. (Trigonella) moorei and T. (Heterotrigona) itama, was studied to describe patterns of resource harvest in disturbed forest areas in Sumatra, Indonesia.

1. Average daily total number of foraging flights per colony was 1200 in T. minangkabau, 2400 in T. moorei and 7000 in T. itama and it was proportional to colony population size. Foragers collecting nectar, pollen or plant resin were respectively 70–80%, 10–20% and <10% in the three species. Pollen was collected most in the morning. Nectar collection peaked in midday in T. itama but continued almost evenly until dusk in T. minangkabau and T. moorei. Resin was collected evenly throughout day.
2. In all the three species the volume of a nectar load carried by a returning forager did not decrease until 1600, followed by slight decline. In the morning the sugar concentration of nectar was almost constantly 30%, and in the afternoon its maximum value reached 60% although some remained 30%. As a result, the mean sugar weight in a nectar load gradually increased until dusk.
3. In T. minangkabau, resin was collected by specialized foragers. Nectar foragers switched to collect pollen and vice versa. There were two types of foraging: “exploitatory” flights, the repitition of short, rather constant flights bringing back full resource loads, and “exploratory” flights, prolonged flight and/or reduction in amount of resources carried. Exploitatory flights followed the exploratory flights which led to discovery of rich sources. Mean duration of exploitatory flights was 7 min during nectar collection, 12 min for pollen and 23 min for resin. Sites of exploited flowers were estimated to be 84–434 m distant from the nest site. Mean duration of flights for nectar collection was 13 min in the morning and 6 min in the late afternoon.

     

Colony development and social structure in a subtropical paper wasp,Ropalidia fasciata (F.) (Hymenoptera: Vespidae)

More than 50% of nests of Ropalidia fasciata were founded by association of foundresses (multifemale nests). The multifemale nests were generally initiated earlier and grew faster than the single-female nests. The survival rate of the multifemale nests was significantly higher than that of single-female nests, and the productivity as measured by the number of cells produced per foundress had a peak at a foundress-group size from 6 to 10. The number of marked foundresses which were seen on their original nest decreased as the colony cycle proceeded, but some of them continued to coexist on the original nests after emergency of many female progeny. Except in the case where a large number of foundresses attended a young nest so that some foundresses could not sit on the nest, the dominance interactions among cofoundresses were mild. More than twothirds (71.4%) of nests (including those at the post-emergence stage) had multiple egg-layers. The foundress association in this species is considered to be beneficial for every foundress because it raises ability to avoid predation or to reconstruct their nests when the nests are destroyed by typhoons.     

Effects of female body size and timing of nest foundation on the reproduction of the Japanese paper wasp,Polistes jadwigae Dalla Torre (Hymenoptera: Vespidae)

Females in hibernation site (FH) were significantly larger than foundresses (FO), suggesting that larger FH were less successful in founding their nest. Body size of FO did not correlate with two nest characters, i.e., the number of cells at pupal stage (CP) and the timing of nest foundation (DP). However, CP negatively correlated with DP in 2 of 3 studied years, suggesting that foundress who founded her nest earlier than others tended to construct larger number of cells irrespective of her body size.     

Evolutionarily stable nesting strategy in a digger wasp.

Two alternative “strategies” will not coexist in a population unless on average they are equally successful. The most likely way for such an equilibrium to be maintained is through something equivalent to frequency-dependent selection. Females of the digger wasp Sphex ichneumoneus (Sphecidae) nest in underground burrows. They usually dig and provision these by themselves but occasionally a nest is jointly occupied. The two wasps fight whenever they meet and in the end only one of the two females lays an egg in the shared nest. Two models based on the theory of mixed evolutionarily stable strategies were developed and tested on comprehensive field data from two North American populations of these wasps. The first model proposes two strategies called founding and joining. Founders start burrows alone, but they are more successful when they are joined by a joiner. At equilibrium founders and joiners are equally successful, which amounts to an amicable, sharing relationship. The predictions of this amicable model are decisively rejected by the data. The second model proposes two strategies called digging and entering. Diggers dig their own burrows but they often have to abandon these burrows because of temporary unsuitability. Enterers move in later, thereby exploiting abandoned burrows as a valuable resource. They do not distinguish an adandoned burrow from one that is still occupied. Therefore sharing of burrows arises as an unfortunate by product of selection for entering abandoned burrows, and Model 2 is not an amicable model. Its quantitative predictions are impressively fulfilled in one population, though not in another population. This is one of the only examples yet known of a mixed evolutionarily stable strategy in nature. Yet the word strategy itself can confuse, and this paper tries the experiment of substituting “decision”, defined as a moment at which the animal commits future time to a course of action.