Social behaviour of two primitively eusocial wasps,Ropalidia sp. nr.variegata andR. gregaria gregaria (Hymenoptera: Vespidae) in the Northern Territory,Australia, with special reference to task specialization and mating inhibition

Social behaviour of females of the Australian paper wasps,Ropalidia sp. nr.variegata (V) andR. gregaria gregaria (G) was observed near Darwin, the Northern Territory, Australia. Both species constructed multiple combs after the emergence of the first progeny. Frequency of intranidal dominance acts in speciesV on the post-emergence nests was significantly higher than that on the pre-emergence nests, while the frequency was constantly high in speciesG throughout the pre- and post- emergence periods. Some females ofG were specialized, at least on single days, to perform the particular task of water collection. In both species the dominant female tended to occupy the largest comb and subordinate females smaller combs. InG, not only the top-ranked female but also low-ranked ones oviposited, and a small proportion of eggs laid by the latter survived at least until the end of observation. InV, each colony had only 1 inseminated female which had highly developed ovaries, suggesting that subordinate females do not mate despite the presence of males. Such a limitation of mating to a single female in a colony has not been described previously in primitively eusocial wasps.     

Comparative reproduction of <Emphasis Type="Italic">Varroa destructor</Emphasis> in different types of Russian and Italian honey bee combs

Earlier studies showed that Russian honey bees support slow growth of varroa mite population. We studied whether or not comb type influenced varroa reproduction in both Russian and Italian honey bees, and whether Russian bees produced comb which inhibited varroa reproduction. The major differences found in this study concerned honey bee type. Overall, the Russian honey bees had lower (2.44 ± 0.18%) levels of varroa infestation than Italian honey bees (7.20 ± 0.60%). This decreased infestation resulted in part from a reduced number of viable female offspring per foundress in the Russian (0.85 ± 0.04 female) compared to the Italian (1.23 ± 0.04 females) honey bee colonies. In addition, there was an effect by the comb built by the Russian honey bee colonies that reduced varroa reproduction. When comparing combs having Russian or Italian colony origins, Russian honey bee colonies had more non-reproducing foundress mites and fewer viable female offspring in Russian honey bee comb. This difference did not occur in Italian colonies. The age of comb in this study had mixed effects. Older comb produced similar responses for six of the seven varroa infestation parameters measured. In colonies of Italian honey bees, the older comb (2001 dark) had fewer (1.13 ± 0.07 females) viable female offspring per foundress than were found in the 2002 new (1.21 ± 0.06 females) and 1980s new (1.36 ± 0.08 females) combs. This difference did not occur with Russian honey bee colonies where the number of viable female offspring was low in all three types of combs. This study suggests that honey bee type largely influences growth of varroa mite population in a colony.     

Acceptance by Honey Bee Guards of Non‐Nestmates is not Increased by Treatment with Nestmate Odours

Honey bee, Apis mellifera, entrance guards use chemical cues to discriminate nestmates from non‐nestmates. Previous research has shown that when wax combs are reciprocally swapped between two colonies, guards become more accepting of workers from the swap partner. However, when combs were transferred only one way, guards in the comb‐receiver colony became more accepting of bees from the comb‐donor colony, but not vice versa. Hence, the increased acceptance of non‐nestmates caused by reciprocal comb swapping was not because of introduced bees acquiring odours from the transferred combs, which was surprising because comb wax was known to affect the odour of bees. In the current experiment, we caused workers to acquire either nestmate or non‐nestmate odours by holding them for 15 min in a tube, which had previously held nestmates or non‐nestmates and then measured their acceptance by entrance guards of nestmate or non‐nestmate hives. When transferred workers had acquired odours of non‐nestmates, acceptance by their own colony’s guards significantly decreased to 66% from 91%. Conversely, the acceptance of non‐nestmates that had acquired odours of the guards’ own nestmates was unchanged, 25% vs. 25%. These results show that when equivalent changes in the odour of introduced bees are made, guards are more sensitive to changes that cause nestmates to acquire non‐nestmate odours than vice versa. These results are also a likely explanation for the earlier and surprising results from the unidirectional comb swap experiment ( Couvillon et al. 2007 ). We make a hypothesis for the underlying mechanism in terms of a multidimensional recognition cue space.     

Evaluation of drone brood removal for management of Varroa destructor (Acari: Varroidae) in colonies of Apis mellifera (Hymenoptera: Apidae) in the northeastern United States

The efficacy of drone brood removal for the management of Varroa destructor Anderson & Trueman in colonies of the honey bee, A. mellifera L., was evaluated. Colonies were treated with CheckMite+ in the fall of 2002. The following spring, quantities of bees and brood were equalized, but colonies were not retreated. The brood nest of each colony consisted of 18 full-depth worker combs and two full-depth drone combs. Each worker comb had <12.9 cm2 of drone cells. Standard management practices were used throughout the season. Colonies were randomly assigned to one of two groups. In the control group, drone combs remained in place throughout the season. In the treatment group, drone combs were removed on 16 June, 16 July, 16 August, and 16 September and replaced with empty drone combs (16 June) or with drone combs removed on the previous replacement date. In the early fall, the average mite-to-bee ratio in the control group was significantly greater than the corresponding ratio in the treatment group. Drone brood removal did not adversely affect colony health as measured by the size of the worker population or by honey production. Fall worker populations were similar in the two groups. Honey production in treatment colonies was greater than or similar to production in control colonies. These data demonstrate that drone brood removal can serve as a valuable component in an integrated pest management program for V. destructor and may reduce the need for other treatments on a colony-by-colony basis.     

The relationship between comb age and performance of honey bee (Apis mellifera) colonies

A study on the relationship between the age of comb and the activity of the hybrid Carniolan honey bee colonies in collecting pollen activity, worker brood production, colony strength, and honey yield was conducted. In comparison to colonies with combs aged 4-years, colonies with combs aged 1, 2 and 3-years significantly exceeded in the number returning workers, number returning workers with pollen loads, rate of storing pollen, rate of worker brood production, and size of colony population. Colonies with combs aged 1, 2 and 3-years produced significantly more honey than colonies with combs aged 4-years (5.25, 4.90 and 4.65 kg/colony vs. 4.45 kg/colony, respectively). It can be concluded that the foraging rate, gathering and storing pollen, brood production, colony population size, and honey yield significantly depended on the age of combs. Beekeepers can replace old combs with new ones to increase brood and honey production.     

Rediscovery of the workerless inquiline antPogonomyrmex colei and additional notes on natural history (Hymenoptera: Formicidae)

Summary Pogonomyrmex colei is a workerless inquiline ant known only from nests ofP. rugosus, its closest relative. Ten of 776 (1.3%) host nests were parasitized at a site in central Arizona, while none of 1499 potential host colonies were parasitized at two other locales. Colonies ofP. colei are perennial, and host alate females in 9 of 10 colonies demonstrates that host queens survive parasitism. Three of 10 colonies died over 19 colony years of observation, while only 1 of 601 colonies became newly parasitized. Mating occurs in morning for up to 2–3 days following summer and fall rains and in afternoon during cool fall days. Mating is intranidal just outside the nest entrance, with males returning to the natal nest. MaleP. colei may be flightless because their wing area is reduced compared to host males. Females fly from the nest and locate potential host colonies by following trunk trails. Workers are the largest barrier to nest establishment, as they removed over 90% ofP. colei females placed in trunk trails or that entered host nests. Males and females ofP. colei andP. anergismus, the only other congeneric inquiline species, are diminutive compared to their hosts, with females 30% lighter than host workers. Fat content is lower and water content is higher inP. colei andP. anergismus females than in their hosts.     

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.     

Honey bees (Apis mellifera) reared in brood combs containing high levels of pesticide residues exhibit increased susceptibility to Nosema (Microsporidia) infection

Nosema ceranae and pesticide exposure can contribute to honey bee health decline. Bees reared from brood comb containing high or low levels of pesticide residues were placed in two common colony environments. One colony was inoculated weekly with N. ceranae spores in sugar syrup and the other colony received sugar syrup only. Worker honey bees were sampled weekly from the treatment and control colonies and analyzed for Nosema spore levels. Regardless of the colony environment (spores+syrup added or syrup only added), a higher proportion of bees reared from the high pesticide residue brood comb became infected with N. ceranae, and at a younger age, compared to those reared in low residue brood combs. These data suggest that developmental exposure to pesticides in brood comb increases the susceptibility of bees to N. ceranae infection.     

Fighting and mating behaviors of dimorphic males in the ant

Colony composition inCardiocondyla wroughtoni and the fighting and mating behaviors of 2 types of males, alates and ergatoids, are described. This species is polygynous, with a mean of 7.0 queens per nest, and forms polycalic colonies. Within nests, ergatoid males fight with each other, leading to the death of all but one in single nests. On the other hand, alate males exhibit no aggressive behavior towards any of their colony members. Both types of males conduct intranidal mating with their sisters, though the alate males also conduct nuptial flights. Many alate females leave their maternal nest even if they have already been inseminated by intranidal mating.     

Reproductive behavior of the territorial herbivoreStegastes nigricans (Pisces: Pomacentridae) in relation to colony formation

Reproductive behavior of the herbivorous damselfishStegastes nigricans was studied. Both males and females held individual feeding territories, which were distributed contiguously and formed colonies. Spawning season was from June to September. Females visited males' territories and spawned eggs on the algal nest. Males tended eggs until hatching but exhibited less care taking behavior than other damselfishes. The mating system was promiscuous in that both males and females spawn with several mates, even within a single day. However, only 1 female was accepted in the nest at a time. The restriction of the number of mates in the nest may be an adptation to prevent the sneaking which was often conducted by other males of the same colony, and may also be avoidance of disturbing the spawning female. Most spawnings were done among members of the same colony and inter-colonial spawnings were rare (27 cases of 333 spawnings observed). The results of this study suggest that colonial distribution ofS. nigricans may bring several benefits in reproduction to the colony members (e.g. easy mate findings within fellow colony members), but also seems to bring costs (e.g. restriction of number of mates).     

Comb cutting inRopalidia plebeiana: a new process of colony fission in social wasps (Hymenoptera: Vespidae)

Summary Females of an Australian polistine wasp,Ropalidia plebeiana, often use their mandibles to cut their nest-comb in spring, dividing it into two or more completely independent nests. Prior to the division, each of the major egg layers, often with some subordinates, tended to occupy a different part of a single comb. These females gnawed cells in the intermediate zone between such territories, and ultimately divided the comb. Many other females also built new nests near the nest aggregations, but addition of new nests by comb cutting represented 34.8 % of the increase in nest number. This method of colony fission is so far unknown in any eusocial Hymenoptera.     

Combs and sexual selection in black grouse (Tetrao tetrix)

We studied supra-orbital combs in lekking black grouse (Tetrao tetrix) in relation to sexual selection at five leks in Finland1991-1998 and four leks in Sweden 1992-1995. Comb size wasestimated in two ways: by observing its natural size in thefield at different behaviors ("observed comb size"), and bymeasuring the comb size from captured birds ("measured combsize"). The size of combs is highly variable, and individualscan change it within seconds. Males express their larger combsduring display, as compared to other behaviors. Observed mean comb sizes were larger on leks with a higher number of malesand a higher number of copulations. Measured and observed combsizes and copulatory success did not significantly correlatewhen all males where analyzed, but a positive and significantrelationship between observed comb size and copulatory success was found within males that achieved copulations. Measured comblength correlated positively with the amount of testosterone.While females were present on the lek, displaying and successfulmales showed the largest observed comb size. When we comparedobserved comb size during fighting between successful and unsuccessfulmales and correlated comb size of pairs of fighting males withtheir fighting activity, no significant differences in combsize were found. The result that comb size correlated significantlywith an increase in testosterone level and that larger combsize, within successful males, predicted higher copulatorysuccess suggests that combs may be a cue for females to assessmale quality. The lack of a significant relationship betweenobserved comb size and fighting behavior suggests that combsize either has minor importance in male-male signaling onthe lek or that males may express similar-sized combs duringfighting to avoid serious fights and thus risk of comb injuries.     

The mite Varroa jacobsoni does not transmit American foulbrood from infected to healthy colonies

The present study was conducted to determine whether Varroa jacobsoni can transmit American foulbrood (AFB), caused by the bacterium Paenibacillus larvae to healthy colonies by the surface transport of spores. Five two-storey Langstroth colonies of Apis mellifera ligustica were infested by placing a sealed brood comb, with 10% Varroa prevalence, between the central brood combs of each colony. Two months later the colonies were inoculated with P. larvae by adding brood comb pieces with clinical signs of AFB (45±5 scales per colony). After 60 days the brood area was completely uncapped by means of dissecting needles and tweezers, separating the Varroa mites from the larvae and the collected mites were introduced at a rate of 51 per colony into four recipient hives placed in an isolated apiary. Twenty female Varroa specimens were separated at random and observed by SEM. Paenibacillus larvae spores were found on the dorsal shield surface and on idiosomal setae. All colonies died after 4–5 months due to a high incidence of varroosis. No clinical AFB symptoms or P. larvae spores were observed in microscopic preparations. It is concluded that Varroa jacobsoni does not transmit AFB from infected to healthy colonies; it does, however transport P. larvae spores on its surface.     

Comb Wax Salvage by the Red Dwarf Honeybee, <Emphasis Type="Italic">Apis florea</Emphasis> F.

Cannibalizing wax from a deserted nest and reusing it in construction of a new nest is only known for absconding colonies of red dwarf honeybee, Apis florea. We tested whether A. florea would preferentially choose to salvage wax from their own, original natal comb over that of other conspecific combs (test 1) and whether they would salvage wax from comb facsimiles of A. florea combs (test 2) fashioned from the combs of A. cerana, A. dorsata, and A. mellifera. In test 1, preferences for natal comb were significantly greater than for non-natal combs. In test 2, no wax was collected from heterospecific combs. It is evident that wax discrimination is context-dependent and that there is considerable genetic variation for the wax-salvaging trait.     

Process of comb division of reused nests in the Australian paper waspRopalidia plebeiana (Hymenoptera,Vespidae)

Summary InRopalidia plebeiana, combs made in the previous year are often reused by foundresses in the following spring and structurally divided into subnests by them. Close observations of the comb dividing process revealed that: 1) combs that had multiple brood areas (areas where cells had eggs) were frequently divided into subnests, while those with single brood areas were never divided; 2) groups of foundresses each occupied a particular brood area even before comb division started; and 3) frequency with which a foundress practiced comb cutting was independent of her social status or oviposition frequency. These observations suggest that the initial grouping of foundresses that colonize an old nest leads to subsequent comb division, rather than the major egg layers cutting combs to safeguard their oviposition territories from their rivals.     

Colony migration in the tropical honey beeApis dorsata F. (Hymenoptera: Apidae)

Summary To obtain insights into the organization and adaptive significance of seasonal migration by colonies of the giant honey bee,Apis dorsata, we monitored the arrivals and departures of colonies in a rain forest habitat in northeastern Thailand, compared patterns of honey bee abundance with other measures of habitat variability, and observed the role of dance communication in organizing the migratory departure of a colony. Colonies arrived in the area during the end of the dry season, reproduced, and then departed early in the rainy season. During the immigration phase, early-arriving colonies stayed only temporarily, as if assessing habitat quality. Colonies departing after a long stay always left barren combs behind, suggesting that they had left in response to deteriorating resource quality. These observations support the idea that migration allows colonies to track seasonally varying resources in different regions. Our observations of a colony preparing for migration revealed that the dance language is involved in organizing the colony's departure, but that dancers signal only the direction to be taken, rather than, as in dances to feeding sites, both the direction and distance of a particular location.     

Nest-founding behavior induced in the first descendants ofPolistes dominulus Christ (Hymenoptera: Vespidae) colonies

Summary First descendants' founding ability was tested inP. dominulus colonies by experimentally removing the original nests and queens 21–24 days after the first emergence. Foundation tests were carried out after three different periods had elapsed after the removal and the results were analyzed with regards to the effects of participation in social tasks, duration of stay within the colony and seasonal factors. 1.Immediately after the removal, the foundations that occurred were performed almost entirely by workers, regardless of the duration of their stay within the colony. The number of foundations depended on the season in which the females had emerged. 2.Two months and four months after the removal, foundations were made by only a small number of females (workers and non-workers). These females, which were therefore able to found a nest the year they were born, were mostly among the very first-born individuals in the colonies. 3.The following year after hibernation: females (workers and non-workers) which had stayed for only a short time within the colony (1–15 days) had the highest survival rates. The foundation rates among the surviving females depended only on seasonal factors, since the foundation rates were higher among the non-workers which had emerged later in the season. The results throw some light on the first descendants' nest-founding potential in colonies developing under natural conditions, where many workers in fact stay for only a short time at the nest.     

Old comb for nesting site recognition by <Emphasis Type="Italic">Apis dorsata?</Emphasis> Field experiments in China

The Asian giant honey bee, Apis dorsata, often conducts seasonal, long-distance migrations in southern China, between a preferred tree (having more than one nest) and alternate sites. Although worker bees cannot make a round-trip journey, colonies re-utilize preferred trees after an absence of several months. We performed comb experiments in which bases and all abandoned combs were entirely scraped off trees and their sites covered with plastic, or comb was moved to trees of the same species. Swarms of giant honey bees investigated trees where combs were removed and continued to nest on the same trees. In contrast, placing combs in nets on previously used trees, or on nearby trees of the same species, did not attract more swarms. The same number of colonies that left them returned to previously occupied trees. Our findings suggest that direct olfactory or sensory contact with old comb bases might regulate nest establishment, but individual trees, lacking normal visual or chemical cues of old nests, are relocated using behavioral devices that remain to be elucidated. Received 12 February 2007; revised 5 June 2007; accepted 13 September 2007.     

Sub-lethal effects of pesticide residues in brood comb on worker honey bee (Apis mellifera) development and longevity

Background

Numerous surveys reveal high levels of pesticide residue contamination in honey bee comb. We conducted studies to examine possible direct and indirect effects of pesticide exposure from contaminated brood comb on developing worker bees and adult worker lifespan.

Methodology/Principal Findings

Worker bees were reared in brood comb containing high levels of known pesticide residues (treatment) or in relatively uncontaminated brood comb (control). Delayed development was observed in bees reared in treatment combs containing high levels of pesticides particularly in the early stages (day 4 and 8) of worker bee development. Adult longevity was reduced by 4 days in bees exposed to pesticide residues in contaminated brood comb during development. Pesticide residue migration from comb containing high pesticide residues caused contamination of control comb after multiple brood cycles and provided insight on how quickly residues move through wax. Higher brood mortality and delayed adult emergence occurred after multiple brood cycles in contaminated control combs. In contrast, survivability increased in bees reared in treatment comb after multiple brood cycles when pesticide residues had been reduced in treatment combs due to residue migration into uncontaminated control combs, supporting comb replacement efforts. Chemical analysis after the experiment confirmed the migration of pesticide residues from treatment combs into previously uncontaminated control comb.

Conclusions/Significance

This study is the first to demonstrate sub-lethal effects on worker honey bees from pesticide residue exposure from contaminated brood comb. Sub-lethal effects, including delayed larval development and adult emergence or shortened adult longevity, can have indirect effects on the colony such as premature shifts in hive roles and foraging activity. In addition, longer development time for bees may provide a reproductive advantage for parasitic Varroa destructor mites. The impact of delayed development in bees on Varroa mite fecundity should be examined further.     

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.