Caste fate conflict in swarm-founding social hymenoptera: an inclusive fitness analysis

A caste system in which females develop into morphologically distinct queens or workers has evolved independently in ants, wasps and bees. Although such reproductive division of labour may benefit the colony it is also a source of conflict because individual immature females can benefit from developing into a queen in order to gain greater direct reproduction. Here we present a formal inclusive fitness analysis of caste fate conflict appropriate for swarm-founding social Hymenoptera. Three major conclusions are reached: (1) when caste is self-determined, many females should selfishly choose to become queens and the resulting depletion of the workforce can substantially reduce colony productivity; (2) greater relatedness among colony members reduces this excess queen production; (3) if workers can prevent excess queen production at low cost by controlled feeding, a transition to nutritional caste determination should occur. These predictions generalize results derived earlier using an allele-frequency model [Behav. Ecol. Sociobiol. (2001) 50: 467] and are supported by observed levels of queen production in various taxa, especially stingless bees, where caste can be either individually or nutritionally controlled.     

The queen is dead—long live the workers: intraspecific parasitism by workers in the stingless bee Melipona scutellaris

Insect societies are well known for their high degree of cooperation, but their colonies can potentially be exploited by reproductive workers who lay unfertilized, male eggs, rather than work for the good of the colony. Recently, it has also been discovered that workers in bumblebees and Asian honeybees can succeed in entering and parasitizing unrelated colonies to produce their own male offspring. The aim of this study was to investigate whether such intraspecific worker parasitism might also occur in stingless bees, another group of highly social bees. Based on a large-scale genetic study of the species Melipona scutellaris , and the genotyping of nearly 600 males from 45 colonies, we show that ∼20% of all males are workers' sons, but that around 80% of these had genotypes that were incompatible with them being the sons of workers of the resident queen. By tracking colonies over multiple generations, we show that these males were not produced by drifted workers, but rather by workers that were the offspring of a previous, superseded queen. This means that uniquely, workers reproductively parasitize the next-generation workforce. Our results are surprising given that most colonies were sampled many months after the previous queen had died and that workers normally only have a life expectancy of ∼30 days. It also implies that reproductive workers greatly outlive all other workers. We explain our results in the context of kin selection theory, and the fact that it pays workers more from exploiting the colony if costs are carried by less related individuals.     

Intraspecific queen parasitism in a highly eusocial bee

Insect societies are well-known for their advanced cooperation, but their colonies are also vulnerable to reproductive parasitism. Here, we present a novel example of an intraspecific social parasitism in a highly eusocial bee, the stingless bee Melipona scutellaris. In particular, we provide genetic evidence which shows that, upon loss of the mother queen, many colonies are invaded by unrelated queens that fly in from unrelated hives nearby. The reasons for the occurrence of this surprising form of social parasitism may be linked to the fact that unlike honeybees, Melipona bees produce new queens in great excess of colony needs, and that this exerts much greater selection on queens to seek alternative reproductive options, such as by taking over other nests. Overall, our results are the first to demonstrate that queens in highly eusocial bees can found colonies not only via supersedure or swarming, but also by infiltrating and taking over other unrelated nests.     

Networks and dominance hierarchies: does interspecific aggression explain flower partitioning among stingless bees?

1. The distribution of consumers among resources (trophic interaction network) may be shaped by asymmetric competition. Dominance hierarchy models predict that asymmetric interference competition leads to a domination of high quality resources by hierarchically superior species. 2. In order to determine the competitive dominance hierarchy and its effect on flower partitioning in a local stingless bee community in Borneo, interspecific aggressions were tested among eight species in arena experiments. 3. All species tested were strongly mutually aggressive in the arena, and the observed interactions were often lethal for one or both opponents. Aggression significantly increased with body size differences between fighting pairs and was asymmetric: larger aggressors were superior over smaller species. Additional aggression tests involved dummies with surface extracts, and results suggest that species‐ and colony‐specific surface profiles are important in triggering the aggressive behaviour. 4. Sixteen stingless bee species were observed foraging on 41 species of flowering plants. The resulting bee–flower interaction network showed a high degree of generalisation (network‐level specialisation H₂’ = 0.11), corresponding to a random, opportunistic distribution of bee species among available flower species. 5. Aggressions on flowers were rare and only occurred at a low level. The dominance hierarchy obtained in the arena experiments did not correlate significantly with plant quality, estimated as the number of flowers per plant or as total bee visitation rate. 6. Our findings suggest that asymmetries in interference competition do not necessarily translate into actual resource partitioning in the context of complex interacting communities.     

Genetic structure of drone congregations of the stingless bee Scaptotrigona mexicana

Drones of stingless bee species often form distinctive congregations of up to several hundred individuals which can persist over considerable periods of time. Here we analyse the genetic structure of three drone congregations of the neotropical stingless bee Scaptotrigona mexicana employing eight microsatellite markers. Two congregations were close to each other (50 m), the third one was located more than 10 km away from them. This spatial pattern was also reflected on the genetic level : the two close congregations did not show any population sub-structuring, whereas the more distant congregation showed a significant population differentiation to both of them. Population subdifferentiation was however low with F _st values (F _st = 0.020 and 0.014) between the distant congregations, suggesting gene flow over larger distances mediated by the drones of S. mexicana. Based on the genotypic data we also estimated the number of colonies contributing drones to the congregations. The two joint congregations consisted of drones originating from 39,6 colonies, while the third congregation was composed of drones from 21,8 colonies, thus proving that congregations of S. mexicana are constituted of unrelated drones of multicolonial origin. Received 23 April 2007; revised 21 September 2007; accepted 2 October 2007.     

Landmark orientation during the approach to the nest in the stingless beeTrigona (Tetragonisca) angustula (Apidae,Meliponinae)

Summary We displaced a small nest box containing stingless bees (Trigona (Tetragonisca)angustula) over distances of up to 1.6 meters in different directions and counted the numbers of returning foragers to measure the effects of this manipulation on the homing ability of bees. Bees find it hard to locate the nest box when it was displaced more than about 1 m backwards, forwards or sideways relative to the direction into which the nest entrance pointed. They do not find the nest when its height above ground is changed. The bees use landmarks in the vicinity of the nest to locate it: When the nest box is displaced and landmark positions are changed so that their angular position at the new nest site is the same as at the normal nest position their homing ability is less impaired than it is without changes in landmark positions. Our results show that the bees do not use the nest box itself as a landmark until they have approached the nest position to within about 1 meter with the aid of surrounding landmarks.     

Cytogenetic characterization of Partamona cupira (Hymenoptera, Apidae) by fluorochromes

Four colonies of the stingless bee Partamona cupira (Hymenoptera: Apidae) were cytogenetically analyzed using conventional staining and the fluorochromes CMA(3) e DAPI. The females have 2n = 34 chromosomes (2K = 32 Mˉ+2 Aˉ). Some females, however, presented an additional large B acrocentric chromosome, to a total of 2n = 35. Chromosome B and the chromosomal pairs 2, 9 and 10 showed CMA (3) (+) bands, indicating an excess of CG base-pairs. A clear association was verified between the P. helleri B chromosome SCAR marker and the presence of a B chromosome in P. cupira. The data obtained suggests that B chromosomes in P. helleri and P. cupira share a common origin.     

Floral resources collected by four native bees species in southern Mexico

Abstract

Melissopalynological analysis of 14 honey samples of Tetragonisca angustula (two), Scaptotrigona mexicana (six), Melipona beecheii (three) and Melipona solani (three) from Soconusco region in Chiapas, Mexico was performed. A total of 79 taxa were identified, being Asteraceae, Euphorbiaceae, Fabaceae, Melostomataceae and Rubiaceae the predominant pollen types. Melissopalynological analysis reported a monofloral composition for Melipona beecheii honeys, where the most predominant pollen type was Fabaceae (54.2%). T. angustula, Scaptotrigona mexicana and Melipona solani honey samples were multifloral and predominant pollen types were Asteraceae, Euphorbiaceae, Fabaceae, Melastomataceae and Solanaceae. The most frequent polliniferous pollen type were Asteraceae, Bernardia interrupta, Euphorbia heterophyla and Miconia. Shannon–Wiener diversity index (H′) for Melipona beecheii 2016 (2.21) and Melipona solani 2017 (1.64) were lower meanwhile honeys of Melipona beecheii 2017, S. mexicana 2017, Melipona solani 2016 and T. angustula 2017 were considered as fairly diverse, because they visited a wider range of plant species, by preferring shrub and wild strata such as: Asteraceae, Euphorbia, Euphorbiaceae, Miconia and Mimosa. Our results showed that the foraging behaviour of these bee species is polylectic, as no taxa was found to be more than 70% of the counted grains. Since the study of the collection of meliponine flower resources in the southern region of Mexico have not been studied yet, it is not possible to analyse the foraging behaviour, the use of resources and the management of these species in this region.     

Effectiveness of recruitment behavior in stingless bees (Apidae,Meliponini)

Summary We examined the ability of stingless bees to recruit nest mates to a food source (i) in group foraging species laying pheromone trails from the food to the nest (Trigona recursa Smith, T. hypogea Silvestri, Scaptotrigona depilis Moure), (ii) in solitary foraging species with possible but still doubtful communication of food location inside the nest (Melipona seminigra Friese, M. favosa orbignyi Guérin), and (iii) in species with a less precise (Nannotrigona testaceicornis Lep., Tetragona clavipes Fab.) or no communication (Frieseomelitta varia Lep.). The bees were allowed to collect food (sugar solution or liver in the necrophageous species) ad libitum and the forager number to accumulate, as it would do under normal unrestrained conditions. The median number of bees collecting differed considerably among the species (1.0–1436.5). It was highest in the species employing scent trails. The time course of recruitment was characteristic for most of the species and largely independent of the number of foragers involved. The two Melipona species recruited other bees significantly faster than T. recursa, S. depilis, and N. testaceicornis during the first 10 to 30 minutes of an experiment. In species laying a scent trail to guide nestmates to a food source the first recruits appeared with a delay of several minutes followed by a quick increase in forager number. The median time required to recruit all foragers available differed among the species between 95.0 and 240.0 min. These differences can at least partly be explained by differences in the recruitment mechanisms and do not simply follow from differences in colony biomass.