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.     

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.     

Colony cycle in the south-eastern coastal populations of Ropalidia plebeiana, the only Ropalidia wasp occurring in temperate Australia

Of the 22 species of Ropalidia occurring in Australia, only R. plebeiana is distributed in temperate areas. This wasp is abundant in the south‐eastern coastal regions of Australia, where it forms huge nest aggregations. Based on our observations in late autumn and early spring, as well as information in the literature, we outline the colony cycle of R. plebeiana in the south‐eastern coastal area of New South Wales. The nesting period lasts for approximately 9.5 months, which is unusually long for temperate polistine wasps. Most foundresses seem to return to their natal nests in early spring (mid‐August to early September) to start their own colonies by reusing pre‐existing nest combs, and the pre‐emergence stage (before the emergence of the first adult brood) lasts for approximately 3.5 months. Reproductives (gynes and males) start to emerge as adults in early March, and the reproductive‐producing period lasts until the end of May. Both the pre‐emergence stage and the reproductive‐producing period are much longer than in other temperate polistine wasps, which is discussed in terms of fluctuating temperature conditions in the area and the formation of nest aggregations, a peculiar habit for temperate populations of R. plebeiana.     

Altruism and relatedness at colony foundation in social insects

Cooperative nest initiation in social insects is most easily explained when cooperating females are relatives, as is common in polistine wasps. However, recent research has revealed that unrelated ant queens also initiate colonies together. Reproductive dominance hierarchies are absent among unrelated foundresses, which contrasts with the rigid dominance hierarchies found among related foundresses. New field studies of joint nest founding among non-relatives show that cooperation is favored where colonies are clumped and brood raiding is common, so that attaining a large worker force quickly is critical to colony survival. These studies enrich our understanding of the role of relatedness in social groups.     

Seasonal change of nest survival and related aspects in an aggregation ofLasioglossum duplex (Dalla torre), a eusocial halictine bee (Hymenoptera: Halictidae)

Nest survival in an aggregation of a eusocial halictine bee, Lasioglossum duplex, was censused through an annual cycle. Out of 2,500 nests marked at the beginning of solitary phase in the spring, only 25.5% attained eusocial phase. But 60.5% of 636 nests attaining eusocial phase successfully produced sexual offspring. This shows that solitary phase is the most vulnerable period in the annual cycle. A very low productivity in eusocial phase in the census year was clarified from examination of 99 nests in the autumn. The number of prospective foundresses surviving to the next year was estimated upon the number of brood cells in examined nests and of old females surviving these nests. Deviation between this estimate and the number of nests made in the next spring was 8.8%, confirming a drop of population size to less than one third in the next year. Some nests solitarily made in the summer by dispersed females were examined. Productivity in such nests was extremely low, hence these nests should contribute virtually nothing to the next generation. Possible factors affecting the low productivity in eusocial phase were enumerated though none of these were supported by concrete evidence. Some considerations were given on the relation between obtained results and colony life cycle in eusocial insects, particularly in halictine bees.     

Colony Development and Serial Polygyny in the Primitively Eusocial Wasp Belonogaster juncea juncea (Vespidae: Polistinae)

In Cameroon, the social ropalidiine wasp Belonogaster juncea juncea mostly selects human constructions for nesting, but nests installed in the shelter of large boulders were also noted. Nests were founded throughout the year by one female (25.5%) or by associations of two to eight females. Among the 49 colonies studied from foundation, 24 produced at least one adult, and only 5 produced males among the eight that succeeded in producing sexual individuals. After the first dominant female disappeared (died or abandoned the nest, from 77 to 196 days after foundation) the remaining females fought among themselves, leading to a drastic decrease in the total number of brood. They completely abandoned 22 of 24 surviving nests, over a mean colony cycle lasting about 7 months. Hence the colony and nesting cycles were congruent, illustrating that determinate nesting corresponds to the majority of the cases in this species. In the other two nests, some females remained on the nest and began a new colonial cycle illustrating semi-independence between colony and nesting cycles. Up to four successive cycles were completed in this way in the same nest, thus illustrating the phenomenon of serial polygyny (intermediate between determinate and indeterminate nesting cycles). Serial polygyny in independent founding polistine wasps, previously documented only once, has implications for studies attempting to identify factors involved in regulating the colony and nesting cycles in tropical social wasps.     

Fungus garden platforms improve hygiene during nest establishment in <Emphasis Type="Italic">Acromyrmex</Emphasis> ants (Hymenoptera,Formicidae, Attini)

Physically isolating organisms from disease agents should reduce the likelihood of disease transmission and infection, and increase survival and growth, particularly in more vulnerable, early ontogenetic stages. During nest founding in fungus-growing ants, foundresses of most genera use a garden platform to isolate the incipient fungal garden from the soil of the underground chamber. We studied nest founding in Acromyrmex octospinosus to test the hypotheses that the use of a platform (rootlets used to suspend the fungus garden): (i) reduces the likelihood that the garden will be contaminated by soil-borne microbial pathogens; (ii) results in more rapid growth of a young colony; and (iii) increases colony survival. We manipulated natural incipient nests to have gardens either in contact with or isolated from soil surrounding the chamber, and nests with and without foundresses present. We found a higher incidence of infection in gardens that were in contact with the chamber soil and without queens, compared with gardens isolated from the chamber soil with and without foundresses. The production of eggs, larvae and pupae, as well as leaf area harvested, were significantly different between nests with and without platforms, but there were no differences in the production of workers nor garden biomass. Likewise, there were no differences between treatments in colony survival rates over 8 weeks. Using smaller incipient gardens, however, gardens with and without platforms differed in survivorship rates after 24 hours. The results indicate that the use of a platform to cultivate an incipient fungal garden is an adaptation to reduce soil-borne diseases and increase colony performance. Received 28 July 2006; revised 15 November 2006; accepted 22 November 2006.     

Philopatry, nest choice, and aggregation temporal–spatial change in the digger wasp Cerceris arenaria (Hymenoptera: Crabronidae)

Studying temporal and spatial changes of aggregations of digger wasps through nesting seasons is interesting because of its link to social evolution via a semi-social pathway, but information about this topic is scarce. An aggregation of Cerceris arenaria Latreille was studied during 1997–1999 and for a short period in 2001. The colony grew in area and number of nests through the first three seasons, but the aggregation reduced in 2001, and this decrease was confirmed by a count of nests made in 2003 and 2005. The positions of the nest entrances remained almost constant over the years, at least in high-nest-density areas, because of the repeated use of the same burrows from which they emerged. This fact permits the population to exploit for many years a restricted area and could act as a nest-density-regulation method for fossorial species that rarely dig new nests, preventing an excessive nest density. The newly emerged females did not dig new nests, showed a high philopatry in the choice of their first nest, and preferred to look for other nests to occupy close to the ones recently abandoned (mainly for an undergone usurpation by a conspecific female). Philopatry, along with lack of new nest digging and position-dependent nest choice, is probably an important factor maintaining spatial stability of the aggregation through the years and could represent a first step in the evolution of a higher sociality in apoid burrowing Hymenoptera.     

Colony Foundation and Nestmate Recognition in the Social Wasp,Belonogaster petiolata

Emergence from hibernation and synchronized nest foundation by foundresses of B. petiolata in the southern Transvaal, appeared to be stimulated largely by a rise in temperature during early and mid August of each season. Initial foundresses on newly built nests were joined by other foundresses during the first few weeks of the nesting season to form foundress associations. Although females were strongly philopatric and able to discriminate between nestmates (from their natal nests) and non-nestmates, former nestmates did not reassociate to a significant degree on new nests. Thus, associated foundresses were, on average, not closely related. Rather than facilitating the reassociation of former nestmates, the advantage of philopatric behaviour appears to lie in ensuring that foundresses return to an area of relatively high nesting density, where their chances of being involved in multiple-foundress groups (irrespective of intra-colonial relatedness) are high. Non-philopatric females which nest away from localities of high nesting density have a low probability of being involved in associations, and may therefore have to attempt colony foundation alone, with little or no chance of reproducing successfully.     

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.     

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.     

Reconstruction ofRopalidia fasciata nests by progeny females after a typhoon and its significance in the social evolution of wasps

A strong typhoon, Typhoon 13, which swept through Okinawa on 2 and 3 September 1993, knocked down 57 to 61 of 97Ropalidia fasciata nests. Thirty-five colonies soon reconstructed their nests, and these nests produced 54±46 adults per nest during the remaining two and a half months before winter. The number of females involved in nest reconstruction was 21.3±15.7 per nest. The frequency of foraging activity was significantly higher in nests being reconstructed than in normal nests. As only one out of 205 foundresses marked in spring was found in early September, most cases of nest reconstruction were considered to be made exclusively by females that emerged during the spring and summer of 1993 (progeny females). All of the reconstructed nests but one produced adult females. Many females (31%) collected from the two reconstructed nests were inseminated. As all the reconstructed nests became vacant by early January, indicating completion of the colony cycle, and a large number of nests were established in the spring of 1994, females emerging from reconstructed nests in 1993 probably become foundresses in 1994.Ropalidia fasciata is considered to be, at least partially, bivoltine in Okinawa. The significance of these facts for the evolution of multi-queen social systems in the Polistinae is discussed.     

Spatial Nest‐Settlement Decisions in Digger Wasps: Conspecifics Matter more than Heterospecifics and Previous Experience

Research into the driving forces behind spatial arrangement of wasp nests has considered abiotic environmental factors, but seldom investigated attraction or repulsion towards conspecifics or heterospecifics. Solitary female digger wasps (Hymenoptera) often nest in dense aggregations, making these insects good models to study this topic. Here, we analysed the nesting patterns in an area shared by three species of the genus Bembix, in a novel study to discover whether female wasps are attracted to or repulsed by conspecific nests, heterospecific nests or their own previously established nests when choosing nest‐digging locations. Early in the season, each species showed a clumping pattern of nests, but later in the season, a random distribution of nests was more common, suggesting an early conspecific attraction. Such behaviour was confirmed by the fact that females started building their nests more frequently where other females of their species were simultaneously digging. The distances between subsequent nests dug by individual females were shorter than those obtained by random simulations. However, this pattern seemed to depend on the tendency to dig close to conspecifics rather than remain in the vicinity of previous nests, suggesting that females' experience matters to future decisions only on a large scale. Nesting patches within nest aggregations largely overlapped between species, but the nests of each species were generally not closer to heterospecific nests than expected by chance, suggesting that females are neither repulsed by, nor attracted to, congenerics within nest aggregations. A role of the spatial distribution of natural enemies on the observed nesting patterns seemed unlikely. Bembix digger wasp nest aggregations seem thus to be primarily the result of female–female attraction during nest‐settlement decisions, in accordance with the ‘copying’ mechanisms suggested for nesting vertebrates.     

Benefits of foundress associations in the paper wasp Polistes dominulus: increased productivity and survival, but no assurance of fitness returns

Successful Polistes dominulus nests can be started by one ormore nest founding queens (foundresses). Consequently, thereis much interest in the specific benefits that induce cooperationamong foundresses. Here, we experimentally demonstrate one majorbenefit of cooperation, namely that multiple foundresses increasecolony productivity. This increase is close to the value predictedby subtracting the productivity of undisturbed single-foundresscolonies from the productivity of undisturbed multiple-foundresscolonies. However, we found no evidence that an associatingfoundress' contribution to colony growth is preserved if shedisappears (assured fitness returns). Our correlational datasuggest that cooperation provides survival benefits, multiple-foundresscolonies are more likely to survive to produce offspring thanare single-foundress colonies, and individual foundresses inmultiple-foundress groups are less likely to disappear beforeworker emergence than foundresses nesting alone. Therefore,association provides substantial productivity and survival benefitsfor cooperating foundresses.     

Variable worker behaviour in the weakly eusocial sweat bee, <Emphasis Type="Italic">Halictus sexcinctus</Emphasis> Fabricius

Summary Studies of eusocial halictines suggest that workers have many reproductive options, including sterile altruism in the maternal nest, combined helping and personal reproduction, and diapause and spring nest founding. How and when workers exercise these various options influences the strength of colony social organization. Halictus sexcinctus exhibits highly polymorphic social behaviour, with solitary colonies in central Europe and both eusocial and communal colonies in southern Greece. Indirect evidence suggests that some worker-brood females are actually gynes. A distinctly bimodal size distribution among foundresses in 1998, the lower size peak being close to the modal body size of workers from 1997, suggests that large worker-brood females overwinter and return to the aggregation as eusocial foundresses. Other first-brood females remain in the maternal nest as workers, although few can be classified as classical, sterile altruists. Only 17% of older, healthy workers are sterile (i.e. had ovarian development scores 0.1), whereas about 83% are reproductive, exhibiting at least one 1/4-developed oocyte. About 57% of older workers have at least one fully or 3/4 developed oocyte, signifying that they are ready or almost ready to lay. Sterile workers exhibit greater total wear (combined mandibular and wing wear) scores than reproductive workers, suggesting that they are older, have higher activity rates, or both.     

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.     

Soil softness, nest initiation and nest sharing in the wasp, Cerceris antipodes (Hymenoptera: Sphecidae)

Abstract. 1. The relationship between soil softness and nest initiation was studied at two aggregations of a ground nesting sphecid wasp, Cerceris antipodes Smith, in New South Wales, Australia.
2. Nests of C.antipodes were initiated throughout the active season. Nests were started more frequently during moist conditions and when more wasps were active.
3. Nest initiation was clumped on days after periods of rain. Nest initiation was only observed when the soil was soft.
4. Two experimental plots that were watered had significantly softer soil and more nests initiated on them than two control plots. Searching female wasps spent considerably more time on the watered plots than the controls.
5. Nest initiation through hard surface soils presents a problem to C.antipodes. Soil softness must be considered in any analysis of nest sharing in this species.     

Comparison of investment in nest construction by the foundresses of consubgenericPolistes wasps,P. (polistes) riparius andP. (P.) chinensis (hymenoptera, vespidae)

Some metric characters of nests built during the founding phase by foundresses were compared between two consubgenericPolistes wasps,P. (Polistes) riparius andP. (P.) chinensis, the former of which inhabit higher latitudes. Volumes and dry weights ofP. riparius nests were strikingly larger than those ofP. chinensis, even when standardized by the foundress weight (2.5 times for volume and 2 times for weight), showing that foundresses ofP. riparius invest much more in the nest construction than those ofP. chinensis. However, percent weights of oral secretion used for nests to total nest weights were smaller inP. riparius than inP. chinensis (52.1% vs. 60.4%). The differences in the investment by foundresses of the two species in the construction and maintenance of nests were discussed in relation to climatic and other environmental factors.     

Water loss in desert ants: caste variation and the effect of cuticle abrasion

Water loss rates (WLRs) varied across castes (workers > alate males > alate females) for desert ants in the genera Aphaenogaster, Messor and Pogonomyrmex. Exposure to soil caused increased WLRs in workers and foundresses (mated, dealate females), apparently because of cuticular abrasion caused by nest excavation. Moreover, field‐collected workers and foundresses from incipient nests (those exposed to soil) had similar WLRs, as did unabraded workers and alate females (those unexposed to soil). These data call into question previous adaptive scenarios for differences in WLRs across ant species and castes. For live alate females, WLRs increased over two stages. The first increase occurred immediately after mating, and the second occurred for foundresses collected 2 days later from incipient nests. By contrast, WLRs of HCN‐killed females were unaffected by mating, but increased significantly for foundresses collected from 2‐day‐old nests.     

Relationships between Nest Architecture and Behavior in <Emphasis Type="Italic">Xylocopa virginica</Emphasis> (Hymenoptera: Apidae)

The large carpenter bees nest in reeds, stems and wood. Many other Hymenoptera, including many wasps and some bees, have strong associations between the physical elements of their nests and behavior. Nests of the North American carpenter bee, Xylocopa virginica, were collected in southern Ontario. Nest architecture elements were examined with respect to their impact on life-history elements. In particular, it was determined that the brood cells are provisioned sequentially both within and among branches. There was also no detectable pattern of sex allocation in broods. Nests with branched architecture contained more foundresses, but these foundresses did not appear to contribute in the construction of larger nests. These findings are discussed with respect to other bees, and to the social structure of Xylocopa virginica.