The role of wax and resin in the nestmate recognition system of a stingless bee, Tetragonisca angustula

Recent research has shown that entrance guards of the stingless bee Tetragonisca angustula make less errors in distinguishing nestmates from non-nestmates than all other bee species studied to date, but how they achieve this is unknown. We performed four experiments to investigate nestmate recognition by entrance guards in T. angustula. We first investigated the effect of colony odours on acceptance. Nestmates that acquired odour from non-nestmate workers were 63% more likely to be rejected while the acceptance rate of non-nestmates treated with nestmate odour increased by only 7%. We further hypothesised that guards standing on the wax entrance tube might use the tube as an odour referent. However, our findings showed that there was no difference in the acceptance of non-nestmates by guards standing on their own colony’s entrance tube versus the non-nestmate’s entrance tube. Moreover, treatment of bees with nestmate and non-nestmate resin or wax had a negative effect on acceptance rates of up to 65%, regardless of the origin of the wax or resin. The role of resin as a source of recognition cues was further investigated by unidirectionally transferring resin stores between colonies. Acceptance rates of nestmates declined by 37% for hives that donated resin, contrasting with resin donor hives where acceptance of non-nestmates increased by 21%. Overall, our results confirm the accuracy of nestmate recognition in T. angustula and reject the hypothesis that this high level of accuracy is due to the use of the wax entrance tubes as a referent for colony odour. Our findings also suggest that odours directly acquired from resin serve no primary function as nestmate recognition cues. The lack of consistency among colonies plus the complex results of the third and fourth experiments highlight the need for further research on the role of nest materials and cuticular profiles in understanding nestmate recognition in T. angustula.     

The role of a scent beacon in the communication of food location by the stingless bee, Melipona panamica

Melipona panamica foragers can deposit a scent beacon that influences the orientation of foragers near a food source. In misdirection experiments, newcomers (bees from the same colony as trained foragers) consistently preferred the feeder at which trained foragers had fed (training feeder) over an identical feeder at which bees had never fed (control feeder) even when the training feeder was placed at a site where experienced foragers had never foraged. Through similar misdirection experiments, the effective radius of the scent beacon was determined to be greater than 6 and less than 12 m. Foragers may deposit this beacon during a sequence of departure behaviors performed at the feeder. Prior to leaving the feeder with a load of sugar solution, bees tended to perform the following sequence of behaviors: (1) spinning, (2) grooming, (3) abdomen dragging, (4) excreting anal droplets, and (5) producing sounds, although not all behaviors were performed prior to each departure or at all sucrose concentrations (0.5–2.5 m). As sucrose concentration increased, the number of newcomers significantly increased, and the number of experienced foragers producing sounds and spinning on the feeder increased. The exact source of the scent beacon remains a mystery. However, three important sources have been excluded. When choosing between identical paired feeders, foragers were not attracted to the feeders with (1) anal droplets, (2) extracts of sucrose solution at which foragers had fed, or (3) mandibular gland extracts. They were indifferent to the first two preparations and exhibited only typical alarm behavior towards the mandibular gland (MG) extract: they oriented towards the feeder with MG extract but consistently landed on the feeder with no MG extract. Other authors have suggested that Melipona foragers deposit anal droplets to attract recruits, however the frequency of anal droplet production and the mass of anal droplets produced by M. panamica foragers are negatively correlated with sucrose concentration. Thus the scent beacon is evidently not deposited with anal droplets, infused into the feeder solution, or produced by mandibular glands. Received: 2 September 1997 / Accepted after revision: 30 January 1998     

Identification of trail pheromone compounds from the labial glands of the stingless bee Geotrigona mombuca

Foragers of several species of stingless bees deposit pheromone spots in the vegetation to guide recruited nestmates to a rich food source. Recent studies have shown that Trigona and Scaptotrigona workers secrete these pheromones from their labial glands. An earlier report stated that species within the genus Geotrigona use citral from their mandibular glands for scent marking. Since convincing experimental proof for this conjecture is lacking, we studied the glandular origin of the trail pheromone of Geotrigona mombuca. In field bioassays, newly recruited bees were diverted by artificial scent trails that branched off from the natural scent trail deposited by their nestmates only when they were baited with extracts from the foragers’ labial glands. Compounds extracted from the mandibular glands, however, did not release trail following behavior. This demonstrates that the trail pheromone of G. mombuca is produced in the labial glands, as in Trigona and Scaptotrigona. Furthermore, in chemical analyses citral was identified exclusively in the foragers’ mandibular glands, which disproves its supposed role as a trail pheromone. The labial glands contained a series of terpene- and wax type esters, with farnesyl butanoate as major constituent. We, therefore, postulate that the trail pheromone of G. mombuca is composed of a blend of esters.     

Potential mechanisms for the communication of height and distance by a stingless bee, Melipona panamica

This study investigates the recruitment communication mechanisms of a stingless bee, Melipona panamica, whose foragers can evidently communicate the three-dimensional location of a good food source. To determine if the bees communicate location information inside or outside the nest, we conducted removal experiments by training marked foragers to one of two identical feeders and then separating these experienced foragers from potential recruits as they left the nest. The feeders were positioned to test the communication of each dimension. The results show that recruits do not simply follow experienced foragers to the food source. Height and distance are communicated within the nest, while direction is communicated outside the nest. We then examined the pulsed sounds produced by recruiting foragers. While unloading food, recruiting foragers produced several short pulses and one or more very long pulses. On average, the longest unloading pulse per performance was 31–50% longer (P ≤ 0.018) for bees foraging on the forest floor than for bees foraging at the top of the forest canopy (40 m high). While dancing, recruiting foragers produced sound pulses whose duration was positively correlated with the distance to the food source (P < 0.001). Dancing recruiters also produced several short sound pulses followed by one or more long pulses. The longest dance pulse per performance was 291 ± 194 ms for a feeder 25 m from the nest and 1858 ± 923 ms for a feeder 360 m away from the nest. The mechanism of directional communication remains a mystery. However, the direction removal experiment demonstrates that newcomers cannot use forager-deposited scent marks for long-distance orientation (>100 m from the nest). Received: 25 September 1997 / Accepted after revision: 31 May 1998     

Antipredation role of clumped nesting by marsh-nesting red-winged blackbirds

Summary Red-winged blackbirds, Agelaius phoeniceus, breed in marshes in high densities and their nests are frequently clumped. Because predation is consistently the most important cause of redwing nesting mortality, high densities of breeding individuals could be an anti-predation adaptation. In our study site predation by marsh wrens, Cistothorus palustris, was the main cause of redwing nesting losses. In situations when marsh wrens were near, predation rates on redwing nests decreased with increasing female density. Group life could reduce predation because of improved nest defense, selfish herd effects, or predator dilution effects. We differentiated between these possibilities by introducing experimental colonies consisting of 3, 6, and 9 artificial nests near and away from active redwing nests. The experimental colonies near active nests suffered less predation, but predation rates were not correlated with colony size or a nest's location within the colony. Therefore, the advantage of group life in this population is probably mutual nest protection.     

Aerial defense of the nest by workers of the stingless bee Trigona (Tetragonisca) angustula (Latreille) (Hymenoptera: Apidae)

Summary The stingless bee Trigona (Tetragonisca) angustula has a sophisticated defense strategy against flying insect predators at the entrance of its nest. Groups of worker bees hover on both sides in front of the nest entrance tube, facing a flight corridor leading to the nest. Intruders which enter this corridor are attacked by these bees from the side and from behind and are forced to the ground by biting bees clinging to their wings. T. angustula is subject to predation by Lestrimelitta limao, a cleptobiotic stingless bee which performs organized raids on other nests to rob food supplies, larval provisions and nest constructing material. The presence of citral, released by L. limao during the raids, leads to a rapid increase in the number of hovering guard bees in front of T. angustula nests. This recruitment in response to citral suggests that the defense behavior in T. angustula has evolved under the pressure of L. limao raids and that citral functions in T. angustula as an alarm kairomone.     

The habitat saturation hypothesis and sociality in an allodapine bee: cooperative nesting is not “making the best of a bad situation”

Several factors thought to be important for the evolution of cooperative breeding in vertebrates have received little attention in facultatively social insects. One of these, the “habitat saturation hypothesis” of Selander (1964), predicts that colony sizes will be greater in breeding units where dispersal opportunities are limited, suggesting that group living is a secondary option to independent reproduction. The Australian allodapine bee Exoneura bicolor exhibits a number of traits that occur in cooperatively breeding bird species, including long life-span, repeated opportunities for reproduction, and vulnerability to brood predation and parasitism. We experimentally examined the effect of a potentially limiting environmental factor, nesting substrate availability, as an agent influencing sociality in E. bicolor. We manipulated nesting substrate availability in two separate locations during a time when foundress dispersal is common. No significant difference was found between colony sizes in cases where dispersal options were abundant and cases where dispersal options were limited. An increase in opportunities for dispersal did not lead to higher rates of independent nesting, suggesting that cooperative nesting is a preferred strategy regardless of distance-related costs of dispersal. Reproductivity per female and brood survival were examined as factors selecting for group living. Low survival of brood in single-female nests has the potential to select for cooperative nesting in this bee. Received: 29 September 1995/Accepted after revision: 24 June 1996     

The role of learning in the mating biology of a sweat bee Lasioglossum zephyrum (Hymenoptera: Halictidae)

Summary Results from experiments on the role of learning in the mating biology of a sweat bee, Lasioglossum zephyrum (Hymenoptera: Halictidae), are described in this paper. Male learning of individual female odors is important in natural populations (Table 1), as has been shown in the laboratory. Four other hypotheses are unlikely to account for the observed behavior: (1) Female odors dissipate rapidly; (2) Males learn and then avoid the study area; (3) Males or females produce repellents which are effective against other males; or (4) males recognize their own odor on previously-contacted females, which they subsequently avoid. Regarding questions of male preference and optimal outbreeding, Tables 2 and 3 show there are no consistent preferences for more novel or less novel female odors.     

Social foraging in stingless bees: how colonies of Melipona fasciata choose among nectar sources

In an experimental set-up, a colony of the stingless bee Melipona fasciata demonstrated its ability to choose the better of two nectar sources. This colony pattern was a result of the following individual behavioural decisions: continue foraging, abandon the feeder, restart foraging and initiate foraging. Only very rarely did individuals switch from one feeder to the other. With the first combination of a rich (2.7 M) and a poor (0.8 M) feeder M. fasciata behaved differently from Apis mellifera. Recruitment occurred to both feeders and the poor feeder was not abandoned completely. When the poor feeder was set to 0.4 M, M. fasciata abandoned the poor feeder rapidly and allocated more foragers to the rich feeder. These patterns were similar to those reported for A. mellifera with the first combination of feeders. Over a sequence of 4 days, experienced bees increasingly determined the colony patterns, and the major function of communication between workers became the reactivation of experienced foragers. The foragers modulated their behaviour not only according to the profitability of the feeder, but also according to previous experience with profitability switches. Thus, experience and communication together regulated colony foraging behaviour. These findings and the results of studies with honeybees suggest that M. fasciata and honeybees use similar decision-making mechanisms and only partly different tools. Received: 21 December 1998 / Accepted: 5 January 1999     

Survival and productivity benefits to social nesting in the sweat bee <Emphasis Type="Italic">Megalopta genalis</Emphasis> (Hymenoptera: Halictidae)

Facultatively solitary and eusocial species allow for direct tests of the benefits of group living. We used the facultatively social sweat bee Megalopta genalis to test several benefits of group living. We surveyed natural nests modified for observation in the field weekly for 5 weeks in 2003. First, we demonstrate that social and solitary nesting are alternative behaviors, rather than different points on one developmental trajectory. Next, we show that solitary nests suffered significantly higher rates of nest failure than did social nests. Nest failure apparently resulted from solitary foundress mortality and subsequent brood orphanage. Social nests had significantly higher productivity, measured as new brood cells provisioned during the study, than did solitary nests. After accounting for nest failures, per capita productivity did not change with group size. Our results support key predictions of Assured Fitness Return models, suggesting such indirect fitness benefits favor eusocial nesting in M. genalis. We compared field collections of natural nests to our observation nest data to show that without accounting for nest failures, M. genalis appear to suffer a per capita productivity decrease with increasing group size. Calculating per capita productivity from collected nests without accounting for the differential probabilities of survival across group sizes leads to an overestimate of solitary nest productivity.     

Inter-litter competition and communal nursing among Norway rats: advantages of birth synchrony

Summary The effect of inter-litter competition on pup survival was investigated in pairs of female rats (Rattus norvegicus) living and breeding in the same environment. If a female gave birth when a 0- to 14-day-old litter was already present in the environment, her pups had a very high chance of surviving, similar to the situation in which no other litter was present. Moreover, the mother was likely to nurse communally with the mother of the 0- to 14-day-old litter. This communal nursing benefitted the newborn pups as evidenced by their being heavier at weaning than litters that were not nursed communally. In contrast, if a female gave birth when a 15-to 28-day-old litter was already present in the environment, her newborn pups were likely to die within 3 days postpartum, owing to the fact that they were often prevented from suckling at their mother's teats, resulting in milk deprivation, and were often beneath the older pups, resulting in physical trauma. These findings suggest that inter-litter competition is an important source of pup mortality when litters are born 15-28 days apart. The data are discussed in terms of the advantages of birth synchrony.     

Reproductive success of solitary and communal Philoponella oweni (Araneae: Uloboridae)

Summary Philoponella oweni (Uloboridae) is a facultatively communal orb-web spider found in arid regions in the southwestern United States. Within one habitat solitary adult females and communal groups of adult females coexist.This study compared the reproductive output (egg production) and reproductive success (production of live offspring) of solitary and communal females. There was no significant difference in the mean number of egg-cases produced by solitary and communal females, but communal females produced significantly more eggs per egg-case. However, communal females suffered significantly higher rates of egg-case parasitism from the pteromalid wasp Arachnopteromalus dasys. As a result, the mean number of liver offspring produced by solitary and communal females did not differ. Thus, both strategies led to roughly equal reproductive success. The advantages of aggregation of adults may be significant in the evolution of social behavior in spiders.     

Effects of aggregation on feeding and survival in a communal wren

Summary Foraging by a social wren, Campylorhynchus nuchalis (Troglodytidae), in a tropical savanna habitat is not enhanced by aggregation. Data for marked individuals show that solitary foraging results in a higher capture rate than foraging near others. We find no evidence of imitative foraging, as individuals actively avoid successful foragers following a capture and successful foragers do not restrict their search to recently productive stations or techniques. Captures are seldom temporally clumped, and clumping is probably not pronounced enough to favor imitation. Juveniles show no greater tendency to respond to captures of others, or to succeed in foraging in a group, than do adults. Aggregation is probably disadvantageous for foraging because of dispersed, scarce, cryptic, and noneruptive prey and because of the searching technique of these foliage-gleaning insectivores. If predator avoidance is enhanced by aggregation, it does not result in either increased survival or increased foraging efficiency in large groups, even by juveniles.     

Ecological costs and benefits of communal behavior in a presocial spider

Summary Philoponella oweni (Uloboridae) is a facultatively communal spider found in the southwestern United States. Solitary females and communal groups of females coexist in the same habitats. Communal females are known to produce more eggs than solitary females. Communal females might acquire the energy for the production of additional eggs by one of three means: they might have a longer life span, and thus more time in which to acquire energy; web maintenance might be more efficient in a communal group; or communal females might spend more time feeding. Field data from a marked population in Arizona showed the third alternative was true.Two possible causes for the greater time spent feeding by communal females were investigated; either the communal web or group is a more efficient trap for insects, or the communal groups are located at sites where insects are more abundant. Analysis of feeding rates and sticky trap data confirmed the second alternative.Potential web attachment sites are limited in some habitats, and mortality during dispersal is very high. When potential web attachment sites are scarce more of the population is found in communal groups; when they are abundant more of the population is solitary. The pattern of dispersal and overwintering is this species suggests that females renewing a colony in the spring could be related.     

Hovering guards of the stingless bee <Emphasis Type="Italic">Tetragonisca angustula</Emphasis> increase colony defensive perimeter as shown by intra- and inter-specific comparisons

Social insects need to defend their nest against robbery, parasitism and predation. The stingless bee Tetragonisca angustula is unique in that it has guards that hover near the nest entrance in addition to guards that stand at the entrance. We tested both the general hypothesis that hovering guards increase the effectiveness with which flying intruders are detected and the specific hypothesis that hovering guards improve the detection of workers of the obligate robber bee, Lestrimellita limao. In an intraspecific study comparing colonies, we found a strong positive relationship between the number of hovering guards and the distance at which a dummy robber bee or L. limao worker, experimentally moved towards the nest entrance, was detected. These results were mirrored in an interspecific study showing that four species of stingless bees with similar population colonies but which lacked hovering guards, detected L. limao only at the nest entrance, in contrast to T. angustula. In addition, we found that a greater number of attacks by guards occurred when dummies were impregnated with citral, a major component of L. limao mandibular gland odour. Our results support the hypothesis that T. angustula hovering guards increase the detection perimeter for flying intruders, especially L. limao.