Nest odor dynamics in the social wasp Vespula vulgaris

We investigated nest odor dynamics in the common yellow jacket, Vespula vulgaris. In six isolated colonies, we tested the aggression rates toward dead nestmates that had been stored for 10 min, 10 and 19 days outside their colonies at –76 °C. The aggression rate increased from about 12% toward recently killed nestmates up to 30% toward nestmates killed 19 days before the experiment. Obviously, the conserved nest odor profile of the nestmates frozen for several days did not match with that of their colony anymore. This indicates a change of the nest odor within the colony. In a second experiment, we kept two colonies each in one nest box with a complete separation of both neighbor nests by a solid wall inside the box for 28 days. In confrontation experiments, the colony members treated dead foragers from the neighbor nest as aggressively as dead foreign, non-neighbor workers (about 39% each) whereas only about 14% reacted aggressively toward dead nestmates. Seventeen days after the replacement of the solid wall by a metallic grid, which allowed no physical contact but air exchange between the two neighbor colonies, the aggression rates toward foreign workers and nestmates remained relatively unaffected whereas it decreased significantly toward dead neighbors to about 11%. These results suggest a nest odor dynamic caused by volatiles transferred between two adjacent colonies, resulting in an equalization of the former colony specific nest odors. A change of nest odor dynamics influenced by volatiles was so far described only for one ant species at all.     

An absence of aggression between non-nestmates in the bull ant <Emphasis Type="Italic">Myrmecia nigriceps</Emphasis>

The ability of social insects to discriminate against non-nestmates is vital for maintaining colony integrity, and in most social insect species, individuals act aggressively towards non-nestmates that intrude into their nest. Our experimental field data revealed that intra-colony aggression in the primitive bulldog ant Myrmecia nigriceps is negligible; our series of bioassays revealed no significant difference in the occurrence of aggression in trials involving workers from the same, a close (less than 300 m) or a far (more than 1.5 km) nest. Further, non-nestmate intruders were able to enter the nest in 60% of our trials; a similar level was observed in trials involving nestmates. These results suggest that workers of M. nigriceps are either unable to recognize alien conspecifics or that the costs of ignoring workers from foreign colonies are sufficiently low to favor low levels of inter-colony aggression in this species.     

Parasitoids and competitors influence colony-level responses in the red imported fire ant,<Emphasis Type="Italic"> Solenopsis invicta</Emphasis>

Social insect colonies respond to challenges set by a variable environment by reallocating work among colony members. In many social insects, such colony-level task allocation strategies are achieved through individual decisions that produce a self-organized adapting group. We investigated colony responses to parasitoids and native ant competitors in the red imported fire ant (Solenopsis invicta). Parasitoid flies affected fire ants by decreasing the proportion of workers engaged in foraging. Competitors also altered colony-level behaviours by reducing the proportion of foraging ants and by increasing the proportion of roaming majors, whose role is colony defence. Interestingly, the presence of both parasitism and competition almost always had similar effects on task allocation in comparison to each of the biotic factors on its own. Thus, our study uniquely demonstrates that the interactive effect of both parasitism and competition is not necessarily additive, implying that these biotic factors alter colony behaviour in distinct ways. More generally, our work demonstrates the importance of studying the dynamics of species interactions in a broader context.     

In-nest environment modulates nestmate recognition in the ant<Emphasis Type="Italic"> Camponotus fellah</Emphasis>

Multiple behavioral and chemical studies indicate that ant nestmate recognition cues are low-volatile substances, in particular hydrocarbons (HCs) located on the cuticular surface. We tested the hypothesis that in the ant Camponotus fellah, nest environment, in particular nest volatile odors, can modulate nestmate-recognition-mediated aggression. Workers were individually confined within their own nest in small cages having either a single mesh (SM = limited physical contact permitted) or a double mesh (DM = exposed to nest volatiles only) screen. Individual workers completely isolated outside their nest (CI) served as control. When reintroduced into a group of 50 nestmates, the CI workers were attacked as alien ants after only 2 weeks of separation, whereas the SM workers were treated as nestmates even after 2 months of separation. Aggression towards DM ants depended on the period of isolation. Only DM workers isolated for over 2 months were aggressed by their nestmates, which did not significantly differ from the CI nestmates. Cuticular HC analyses revealed that the profile of the non-isolated ants (NI) was clearly distinct from that of CI, SM and DM ants. Profile differences matched the aggressive response in the case of CI ants but were uncorrelated in the case of SM or DM ants. This suggests that keeping the ants within the nest environment affected nestmate recognition in additional ways than merely altering their HC profile. Nest environment thus appears to affect label–template mismatch by modulating aggressive behavior, as well as the direction at which cuticular HCs diverged during the separation period.     

Nest- and colony-mate recognition in polydomous colonies of meat ants (Iridomyrmex purpureus)

Workers of polydomous colonies of social insects must recognize not only colony-mates residing in the same nest but also those living in other nests. We investigated the impact of a decentralized colony structure on colony- and nestmate recognition in the polydomous Australian meat ant (Iridomyrmex purpureus). Field experiments showed that ants of colonies with many nests were less aggressive toward alien conspecifics than those of colonies with few nests. In addition, while meat ants were almost never aggressive toward nestmates, they were frequently aggressive when confronted with an individual from a different nest within the same colony. Our chemical analysis of the cuticular hydrocarbons of workers using a novel comprehensive two-dimensional gas chromatography technique that increases the number of quantifiable compounds revealed both colony- and nest-specific patterns. Combined, these data indicate an incomplete transfer of colony odor between the nests of polydomous meat ant colonies.     

Pesticide exposure in honey bees results in increased levels of the gut pathogen <Emphasis Type="Italic">Nosema</Emphasis>

Global pollinator declines have been attributed to habitat destruction, pesticide use, and climate change or some combination of these factors, and managed honey bees, Apis mellifera, are part of worldwide pollinator declines. Here we exposed honey bee colonies during three brood generations to sub-lethal doses of a widely used pesticide, imidacloprid, and then subsequently challenged newly emerged bees with the gut parasite, Nosema spp. The pesticide dosages used were below levels demonstrated to cause effects on longevity or foraging in adult honey bees. Nosema infections increased significantly in the bees from pesticide-treated hives when compared to bees from control hives demonstrating an indirect effect of pesticides on pathogen growth in honey bees. We clearly demonstrate an increase in pathogen growth within individual bees reared in colonies exposed to one of the most widely used pesticides worldwide, imidacloprid, at below levels considered harmful to bees. The finding that individual bees with undetectable levels of the target pesticide, after being reared in a sub-lethal pesticide environment within the colony, had higher Nosema is significant. Interactions between pesticides and pathogens could be a major contributor to increased mortality of honey bee colonies, including colony collapse disorder, and other pollinator declines worldwide.     

Does the queen win it all? Queen–worker conflict over male production in the bumblebee,<Emphasis Type="Italic"> Bombus terrestris</Emphasis>

Social insects provide a useful model for studying the evolutionary balance between cooperation and conflict linked to genetic structure. We investigated the outcome of this conflict in the bumblebee, Bombus terrestris, whose annual colony life cycle is characterized by overt competition over male production. We established artificial colonies composed of a queen and unrelated workers by daily exchange of callow workers between colony pairs of distinct genetic make-up. Using microsatellite analysis, this procedure allowed an exact calculation of the proportion of worker-derived males. The development and social behavior of these artificial colonies were similar to those of normal colonies. Despite a high worker reproduction attempt (63.8% of workers had developed ovaries and 38.4% were egg-layers), we found that on average 95% of the males produced during the competition phase (CPh) were queen-derived. However, in four colonies, queen death resulted in a considerable amount of worker-derived male production. The different putative ultimate causes of this efficient control by the queen are discussed, and we suggest a possible scenario of an evolutionary arms race that may occur between these two female castes.     

Queen movement during colony emigration in the facultatively polygynous ant Pachycondyla obscuricornis

In ants, nest relocations are frequent but nevertheless perilous, especially for the reproductive caste. During emigrations, queens are exposed to predation and face the risk of becoming lost. Therefore the optimal strategy should be to move the queen(s) swiftly to a better location, while maintaining maximum worker protection at all times in the new and old nests. The timing of that event is a crucial strategic issue for the colony and may depend on queen number. In monogynous colonies, the queen is vital for colony survival, whereas in polygynous colonies a queen is less essential, if not dispensable. We tested the null hypothesis that queen movement occurs at random within the sequence of emigration events in both monogynous and polygynous colonies of the ponerine ant Pachycondyla obscuricornis. Our study, based on 16 monogynous and 16 polygynous colony emigrations, demonstrates for the first time that regardless of the number of queens per colony, the emigration serial number of a queen occurs in the middle of all emigration events and adult ant emigration events, but not during brood transport events. It therefore appears that the number of workers in both nests plays an essential role in the timing of queen movement. Our results correspond to a robust colony-level strategy since queen emigration is related neither to colony size nor to queen number. Such an optimal strategy is characteristic of ant societies working as highly integrated units and represents a new instance of group-level adaptive behaviors in social insect colonies.     

Predatory spider mimics acquire colony-specific cuticular hydrocarbons from their ant model prey

The integrity of social insect colonies is maintained by members recognising and responding to the chemical cues present on the cuticle of any intruder. Nevertheless, myrmecophiles use chemical mimicry to gain access to these nests, and their mimetic signals may be acquired through biosynthesis or through contact with the hosts or their nest material. The cuticular hydrocarbon profile of the myrmecophilous salticid spider Cosmophasis bitaeniata closely resembles that of its host ant Oecophylla smaragdina. Here, we show that the chemical resemblance of the spider does not arise through physical contact with the adult ants, but instead the spider acquires the cuticular hydrocarbons by eating the ant larvae. More significantly, we show that the variation in the cuticular hydrocarbon profiles of the spider depends upon the colony of origin of the ant larvae prey, rather than the parentage of the spider.     

Social Hackers: Integration in the Host Chemical Recognition System by a Paper Wasp Social Parasite

 Obligate social parasites in the social insects have lost the worker caste and the ability to establish nests. As a result, parasites must usurp a host nest, overcome the host recognition system, and depend on the host workers to rear their offspring. We analysed cuticular hydrocarbon profiles of live parasite females of the paper wasp social parasite Polistes sulcifer before and after usurpation of host nests, using the non-destructive technique of solid-phase micro-extraction. Our results reveal that hydrocarbon profiles of parasites change after usurpation of host nests to match the cuticular profile of the host species. Chemical evidence further shows that the parasite queen changes the odour of the nest by the addition of a parasite-specific hydrocarbon. We discuss the possible role of this in the recognition and acceptance of the parasite and its offspring in the host colony. Received: 18 November 1999 / Accepted in revised form: 22 December 1999     

The cues have it; nest-based, cue-mediated recruitment to carbohydrate resources in a swarm-founding social wasp

This study explores whether or not foragers of the Neotropical swarm-founding wasp Polybia occidentalis use nest-based recruitment to direct colony mates to carbohydrate resources. Recruitment allows social insect colonies to rapidly exploit ephemeral resources, an ability especially advantageous to species such as P. occidentalis, which store nectar and prey in their nests. Although recruitment is often defined as being strictly signal mediated, it can also occur via cue-mediated information transfer. Previous studies indicated that P. occidentalis employs local enhancement, a type of cue-mediated recruitment in which the presence of conspecifics at a site attracts foragers. This recruitment is resource-based, and as such, is a blunt recruitment tool, which does not exclude non-colony mates. We therefore investigated whether P. occidentalis also employs a form of nest-based recruitment. A scented sucrose solution was applied directly to the nest. This mimicked a scented carbohydrate resource brought back by employed foragers, but, as foragers were not allowed to return to the nest with the resource, there was no possibility for on-nest recruitment behavior. Foragers were offered two dishes—one containing the test scent and the other an alternate scent. Foragers chose the test scent more often, signifying that its presence in the nest induces naïve foragers to search for it off-nest. P. occidentalis, therefore, employs a form of nest-based recruitment to carbohydrate resources that is mediated by a cue, the presence of a scented resource in the nest.     

Clonal structure affects the assembling behavior in the Japanese queenless ant <Emphasis Type="Italic">Pristomyrmex punctatus</Emphasis>

The queenless ant Pristomyrmex punctatus (Hymenoptera: Myrmicinae) has a unique society that differs from those of other typical ants. This species does not have a queen, and the workers lay eggs and produce their clones parthenogenetically. However, a colony of these ants does not always comprise members derived from a single clonal line. In this study, we examined whether P. punctatus changes its “assembling behavior” based on colony genetic structure. We prepared two subcolonies—a larger one comprising 200 individuals and a smaller one comprising 100 individuals; these subcolonies were established from a single stock colony. We investigated whether these subcolonies assemble into a single nest. The genetically monomorphic subcolonies (single clonal line) always fused into a single nest; however, the genetically polymorphic subcolonies (multiple clonal lines) did not tend to form a single colony. The present study is the first to demonstrate that the colony genetic structure significantly affects social viscosity in social insects.     

Out with the garbage: the parasitic strategy of the mantisfly Plega hagenella mass-infesting colonies of the eusocial bee Melipona subnitida in northeastern Brazil

Between April and June of 2012 mantisflies (Plega hagenella) were found to be extensively parasitizing the nests of two groups of managed colonzies of eusocial stingless bees (Melipona subnitida) in the semi-arid region of northeastern Brazil. The mantisfly larvae developed inside closed brood cells of the bee comb, where each mantispid larva fed on the bee larva or pupa present in a single brood cell. Mature mantispid larvae pupated inside silken cocoons spun in place within their hosts' brood cells then emerged as pharate adults inside the bee colony. Pharate adults were never attacked and killed by host colony workers. Instead, colony workers picked up the pharates and removed them from the nest unharmed, treating them similar to the way that the general refuse is removed from the nest. Adult mantispids subsequently eclosed from their pupal exuviae outside the nest. Manipulative experiments showed that post-eclosion adult mantispids placed back within active bee colonies were quickly attacked and killed. These observations demonstrate that pharate and post-eclosion adults of P. hagenella are perceived differently by colony workers and that delayed adult eclosion is an important functional element in the parasitic life strategy of P. hagenella, allowing adults to escape without injury from the bee colonies they parasitize.     

Communal nesting and kinship in degus (<Emphasis Type="Italic">Octodon degus</Emphasis>)

Communal nesting is a fundamental component of many animal societies. Because the fitness consequences of this behavior vary with the relatedness among nest mates, understanding the kin structure of communally nesting groups is critical to understanding why such groups form. Observations of captive degus (Octodon degus) indicate that multiple females nest together, even when supplied with several nest boxes. To determine whether free-living degus also engage in communal nesting, we used radiotelemetry to monitor spatial relationships among adult females in a population of O. degus in central Chile. These analyses revealed that females formed stable associations of > 2–4 individuals, all of whom shared the same nest site at night. During the daytime, spatial overlap and frequency of social interactions were greatest among co-nesting females, suggesting that nesting associations represent distinct social units. To assess kinship among co-nesting females, we examined genotypic variation in our study animals at six microsatellite loci. These analyses indicated that mean pairwise relatedness among members of a nesting association (r=0.25) was significantly greater than that among randomly selected females (r=–0.03). Thus, communally nesting groups of degus are composed of female kin, making it possible for indirect as well as direct fitness benefits to contribute to sociality in this species.     

Temporal genetic structure of a drone congregation area of the giant Asian honeybee (Apis dorsata)

The giant Asian honeybee (Apis dorsata), like all other members of the genus Apis, has a complex mating system in which the queens and males (drones) mate at spatially defined drone congregation areas (DCAs). Here, we studied the temporal genetic structure of a DCA of A. dorsata over an 8-day time window by the genotyping of sampled drones with microsatellite markers. Analysis of the genotypic data revealed a significant genetic differentiation between 3 sampling days and indicated that the DCA was used by at least two subpopulations at all days in varying proportions. The estimation of the number of colonies which used the DCA ranged between 20 and 40 colonies per subpopulation, depending on the estimation procedure and population. The overall effective population size was estimated as high as N _e=140. The DCA seems to counteract known tendencies of A. dorsata for inbreeding within colony aggregations by facilitating gene flow among subpopulations and increasing the effective population size.     

Garden sharing and garden stealing in fungus-growing ants

 Fungi cultivated by fungus-growing ants (Attini: Formicidae) are passed on between generations by transfer from maternal to offspring nest (vertical transmission within ant species). However, recent phylogenetic analyses revealed that cultivars are occasionally also transferred between attine species. The reasons for such lateral cultivar transfers are unknown. To investigate whether garden loss may induce ants to obtain a replacement cultivar from a neighboring colony (lateral cultivar transfer), pairs of queenright colonies of two Cyphomyrmex species were set up in two conjoined chambers; the garden of one colony was then removed to simulate the total crop loss that occurs naturally when pathogens devastate gardens. Garden-deprived colonies regained cultivars through one of three mechanisms: joining of a neighboring colony and cooperation in a common garden; stealing of a neighbor's garden; or aggressive usurpation of a neighbor's garden. Because pathogens frequently devastate attine gardens under natural conditions, garden joining, stealing and usurpation emerge as critical behavioral adaptations to survive garden catastrophes. Received: 16 June 2000 / Accepted in revised form: 14 September 2000     

成都平原区水稻土有机碳剖面分布特征及影响因素

了解环境因素对土壤有机碳剖面分布的影响,是准确拟合土壤有机碳在垂直方向上的连续变化、开展其三维空间分布模拟和估算区域土壤有机碳储量的基础.基于171个土壤剖面采样数据,利用指数递减函数拟合土壤有机碳剖面分布,分析成都平原区水稻土有机碳剖面分布的空间变异特征,并探讨成土母质、土壤类型(亚类和土属)、海拔、与河流距离和土地利用方式对其空间变异的影响作用,进而揭示环境因素对土壤有机碳剖面分布的影响.结果表明,0~20、20~40、40~60和60~100 cm土壤有机碳均值分别为19.42、9.59、5.99和5.20 g·kg-1,表现出显著的剖面递减趋势.土壤有机碳含量主要集中在40 cm以上,占整个土壤剖面的质量分数为72.17%,是研究成都平原区水稻土碳源/汇的关键部分.拟合土壤有机碳剖面分布的指数递减函数的2类参数呈现出一致的空间分布格局,具有空间相关性;参数C和k的块金系数分别为55.400%和47.671%,表明成都平原区水稻土壤有机碳剖面分布受结构性因素和随机性因素共同影响.回归分析揭示,成土母质和土属是影响研究区土壤有机碳剖面分布的主控因素,但海拔、与河流距离和土地利用的作用不容忽视.在拟合成都平原区水稻土有机碳剖面分布、构建其三维空间预测模型和估算土壤有机碳储量时,应重点考虑成土母质和土属的作用.     

High precision during food recruitment of experienced (reactivated) foragers in the stingless bee<Emphasis Type="Italic"> Scaptotrigona mexicana</Emphasis> (Apidae,Meliponini)

Several studies have examined the existence of recruitment communication mechanisms in stingless bees. However, the spatial accuracy of location-specific recruitment has not been examined. Moreover, the location-specific recruitment of reactivated foragers, i.e., foragers that have previously experienced the same food source at a different location and time, has not been explicitly examined. However, such foragers may also play a significant role in colony foraging, particularly in small colonies. Here we report that reactivated Scaptotrigona mexicana foragers can recruit with high precision to a specific food location. The recruitment precision of reactivated foragers was evaluated by placing control feeders to the left and the right of the training feeder (direction-precision tests) and between the nest and the training feeder and beyond it (distance-precision tests). Reactivated foragers arrived at the correct location with high precision: 98.44% arrived at the training feeder in the direction trials (five-feeder fan-shaped array, accuracy of at least ±6° of azimuth at 50 m from the nest), and 88.62% arrived at the training feeder in the distance trials (five-feeder linear array, accuracy of at least ±5 m or ±10% at 50 m from the nest). Thus, S. mexicana reactivated foragers can find the indicated food source at a specific distance and direction with high precision, higher than that shown by honeybees, Apis mellifera, which do not communicate food location at such close distances to the nest.     

Caps and gaps: a computer model for studies on brood incubation strategies in honeybees (<Emphasis Type="Italic">Apis mellifera carnica</Emphasis>)

In addition to heat production on the comb surface, honeybee workers frequently visit open cells (“gaps”) that are scattered throughout the sealed brood area, and enter them to incubate adjacent brood cells. We examined the efficiency of this heating strategy under different environmental conditions and for gap proportions from 0 to 50%. For gap proportions from 4 to 10%, which are common to healthy colonies, we find a significant reduction in the incubation time per brood cell to maintain the correct temperature. The savings make up 18 to 37% of the time, which would be required for this task in completely sealed brood areas without any gaps. For unnatural high proportions of gaps (>20%), which may be the result of inbreeding or indicate a poor condition of the colony, brood nest thermoregulation becomes less efficient, and the incubation time per brood cell has to increase to maintain breeding temperature. Although the presence of gaps is not essential to maintain an optimal brood nest temperature, a small number of gaps make heating more economical by reducing the time and energy that must be spent on this vital task. As the benefit depends on the availability, spatial distribution and usage of gaps by the bees, further studies need to show the extent to which these results apply to real colonies. M. Fehler and M. Kleinhenz contributed equally to this work.     

The alternative Pharaoh approach: stingless bees mummify beetle parasites alive

Workers from social insect colonies use different defence strategies to combat invaders. Nevertheless, some parasitic species are able to bypass colony defences. In particular, some beetle nest invaders cannot be killed or removed by workers of social bees, thus creating the need for alternative social defence strategies to ensure colony survival. Here we show, using diagnostic radioentomology, that stingless bee workers (Trigona carbonaria) immediately mummify invading adult small hive beetles (Aethina tumida) alive by coating them with a mixture of resin, wax and mud, thereby preventing severe damage to the colony. In sharp contrast to the responses of honeybee and bumblebee colonies, the rapid live mummification strategy of T. carbonaria effectively prevents beetle advancements and removes their ability to reproduce. The convergent evolution of mummification in stingless bees and encapsulation in honeybees is another striking example of co-evolution between insect societies and their parasites.