Vera Lucia Imperatriz-Fonseca

Mate number, kin selection and social conflicts in stingless bees and honeybees

Microsatellite genotyping of workers from 13 species (ten genera) of stingless bees shows that genetic relatedness is very high. Workers are usually daughters of a single, singly mated queen. This observation, coupled with the multiple mating of honeybee queens, permits kin selection theory to account for many differences in the social biology of the two taxa. First, in contrast to honeybees, where workers are predicted to and do police each others male production, stingless bee workers are predicted to compete directly with the queen for rights to produce males. This leads to behavioural and reproductive conflict during oviposition. Second, the risk that a daughter queen will attack the mother queen is higher in honeybees, as is the cost of such an attack to workers. This explains why stingless bees commonly have virgin queens in the nest, but honeybees do not. It also explains why in honeybees the mother queen leaves to found a new nest, while in stingless bees it is the daughter queen who leaves.

Diversity, threats and conservation of native bees in the Neotropics

The Neotropics bee fauna is very rich with 5000 recognised species, including 33 genera (391 species) of Meliponini, but it is estimated to be at least three fold greater in species richness. Deforestation, agriculture intensification and introduction/spread of exotic competing bee species are considered the main threats to most indigenous species, although other less obvious causes can affect the populations of some bee species locally. Efforts to conserve the native bee fauna include better knowledge of bee richness and diversity (standardized surveys, larger bee collections and appropriate identification of bee species) and of their population dynamics, raising of public and policy makers’ awareness, commercial applications of bee products and services such as pollination and preservation of natural habitat.ZusammenfassungDie vorliegende Arbeit soll einen Überblick geben über die Diversität und den Artenreichtum der neotropischen Bienenfauna, die Bedrohungen, denen sie ausgesetzt ist, als auch eine Darstellung der Initiativen und Probleme im Artenschutz der einheimischen Bienen. Die Daten beruhen auf umfassenden Literaturrecherchen. Diese Informationen wurden dann von den Autoren diskutiert und in Form relevanter Punkte inhaltlich zusammengefasst. Das Ergebnis zeigt, dass die Neotropis eine artenreiche Bienenfauna aufweist und dass diese sogar unterschätzt wird. Den existierenden 5000 gültigen Artennamen stehen Schätzungen gegenüber, dass diese weniger als ein Drittel der tatsächlich vorkommenden Arten umfassen (Tab. I). Bedrohungen, denen einheimische Bienen der Neotropis ausgesetzt sind, liegen vor allem menschliche Aktivitäten zugrunde, die in drei Kategorien zusammengefasst werden können: Entwaldung, Intensivierung der Landwirtschaft und Einführung fremder Arten. Die Hauptursachen der Entwaldung sind Holzeinschlag, das Sammeln von Feuerholz, die Produktion von Holzkohle und Rodungen zur Schaffung von landwirtschaftlichen und Weideflächen. Der Amazonasregenwald, Mexiko und Zentralamerika weisen die höchsten Entwaldungsraten in Amerika auf, aber auch in den Chaco-Wälder der argentinischen und kolumbianischen Anden schreitet die Entwaldung fort (Tab. II). Die Ausbreitung und Intensivierung der Landwirtschaft wird vielfach als die wichtigste Bedrohung für Bienenarten angesehen. Sie führt zu einer Verringerung der Artendiversität bei Tieren und Pflanzen, verringert das Angebot an Nistmöglichkeiten und Futterquellen, und durch das Ausbringen von Pestiziden und Pflügen der Böden werden sowohl Bienenlarven als auch Adulte getötet. Ironischerweise sind die meisten Kulturpflanzen mehr oder weniger stark auf die Präsenz biotischer Bestäuber angewiesen, wobei die Bienen die wichtigste Gruppe darstellen. Auch die Einführung fremder Bienenarten und anderer exotischer Organismen, die mit der lokalen Bienenfauna in Wechselwirkung treten, kann die einheimische Bienenfauna beeinträchtigen. Die Einführung von Apis mellifera in die Neue Welt zur Steigerung der Honigproduktion und die von exotischen Hummelarten für Bestäubungszwecke hat zu Bedenken Anlass gegeben über die Konkurrenz mit einheimischen Bienen um Futter- und Nistmöglichkeiten, sowie zur Ausbreitung von Krankheiten und Parasiten und zur Hybridisierung mit einheimischen Hummmelarten. Anderen Bedrohungen liegen Trockenzeiten, Überschwemmungen, grossflächige Buschbrände, Hurrikane und die Kontaminierung der Ökosysteme mit Schwermetallen zugrunde. Die Hauptprobleme, denen sich Initiativen zum Artenschutz einheimischer Bienen gegenübersehen, sind fehlende Kenntnisse über Artenreichtum, Diversität, Taxonomie, Populationsdynamik und den Einfluss menschlicher Aktivitäten auf die meisten Bienenarten. Um zu besseren Kenntnissen über Artenreichtum, Diversität und Populationsdynamik zu kommen, ist Öffentlichkeitsarbeit und Aufklärung bei Politikverantwortlichen erforderlich. Hierin können die Kommerzialisierung von Bienenprodukten, sowie Aufklärung über die Bedeutung von Bestäubern und der Schutz natürlicher Habitate eine wichtige Rolle spielen. Bestäuberinitiativen erweisen sich hierbei als wichtige Werkzeuge, um Politiker, die Öffenlichkeit und Forscher in koordinierter Weise zusammenzubringen, Wissen über wichtige Fragen zu schaffen und insbesondere die negativen Auswirkungen bienenbedrohender Aktivitäten in Lateinamerika abzumildern.

Caste Development, Reproductive Strategies, and Control of Fertility in Honey Bees and Stingless Bees

In the highly eusocial honey bees (Apinae) and stingless bees (Meliponinae) marked caste syndromes have evolved. The typical queen and worker characteristics are different from one another not just with regard to the reproductive organs but include many other morphological, physiological, and behavioral differences as well. These striking caste distinctions are the result of postembryonic divergences in development which depend on ecological conditions, mainly on the external factor of larval nutrition. Caste development in social insects in general belongs to the widespread phenomena of insect polymorphism and, in accordance with environmental control, castes in bees were called ecomorphs (de Wilde and Wirtz 1974; de Wilde 1975). The trophogenic basis for caste induction is provided by the nursing workers, although in a colony brood rearing including gyne production is strongly influenced by the queen. The queen also rigidly controls all reproduction of the workers. On the other hand, her own reproductive output depends completely on the helper functions of the workers, her daughters.

Safeguarding pollinators and their values to human well-being

Wild and managed pollinators provide a wide range of benefits to society in terms of contributions to food security, farmer and beekeeper livelihoods, social and cultural values, as well as the maintenance of wider biodiversity and ecosystem stability. Pollinators face numerous threats, including changes in land-use and management intensity, climate change, pesticides and genetically modified crops, pollinator management and pathogens, and invasive alien species. There are well-documented declines in some wild and managed pollinators in several regions of the world. However, many effective policy and management responses can be implemented to safeguard pollinators and sustain pollination services.

Male production in stingless bees: variable outcomes of queen—worker conflict

The genetic structure of social insect colonies is predicted to affect the balance between cooperation and conflict. Stingless bees are of special interest in this respect because they are singly mated relatives of the multiply mated honeybees. Multiple mating is predicted to lead to workers policing each others’ male production with the result that virtually all males are produced by the queen, and this prediction is borne out in honey bees. Single mating by the queen, as in stingless bees, causes workers to be more related to each others’ sons than to the queens sons, so they should not police each other. We used microsatellite markers to confirm single mating in eight species of stingless bees and then tested the prediction that workers would produce males. Using a likelihood method, we found some worker male production in six of the eight species, although queens produced some males in all of them. Thus the predicted contrast with honeybees is observed, but not perfectly, perhaps because workers either lack complete control or because of costs of conflict. The data are consistent with the view that there is ongoing conflict over male production. Our method of estimating worker male production appears to be more accurate than exclusion, which sometimes underestimates the proportion of males that are worker produced.

Biology of the stingless bee Plebeia remota (Holmberg): observations and evolutionary implications

SummaryThe genusPlebeia has a special significance for the study of social evolution of stingless bees: morphologically primitive, its species display a wealth of behavioural evolution, especially with respect to the oviposition process. We comparePlebeia remota with the few other members of the genus studied so far.Related to its subtropical geographical range, brood production is seasonal (there is no brood in the colony in colder months), and adult workers occur as summer and winter bees. The nest is in tree cavities, and the involucrum is absent or restricted to the winter period. Brood cells are arranged in horizontal combs, and new cells are built completely synchronously. Each series consists of up to 50 cells, their number being mainly dependent on colony size. Construction speed is remarkably constant, allowing 4–6 batches per 24 hours. Cell building and provisioning are activities of a small group of specialized workers.The oviposition cycle follows the classical subdivisions for stingless bees. During the patrolling phase a worker may “offer” a trophic egg in a most remarkable way: while retreating backward from the queen she bends the abdomen under thorax and head, and lays an egg on the comb. This egg is eaten by the queen or a worker.The provisioning, oviposition and operculation of all cells occur simultaneously, each cell is provisioned by 4–9 workers. Localization of a cell by the queen may be facilitated by its characteristic guard, which “defends” the cell against the approaching queen. The degree of synchronization within a batch is very high: the duration per cell lasts 420–950 sec, the batch of up till 50 cells needs only 557–1160 sec. Operculation is done by a worker that was not involved in the previous steps.Males are generally produced by the queen. Several male producing cycles per year occur. In orphan colonies laying workers give rise to males, and in queenright colonies workers may occasionally reproduce as well.Division of labour follows the general pattern for stingless bees; however, cell building and provisioning are activities of a specialized group of workers.

Olfactory eavesdropping by a competitively foraging stingless bee, Trigona spinipes.

Signals that are perceived over long distances or leave extended spatial traces are subject to eavesdropping. Eavesdropping has therefore acted as a selective pressure in the evolution of diverse animal communication systems, perhaps even in the evolution of functionally referential communication. Early work suggested that some species of stingless bees (Hymenoptera, Apidae, Meliponini) may use interceptive olfactory eavesdropping to discover food sources being exploited by competitors, but it is not clear if any stingless bee can be attracted to the odour marks deposited by an interspecific competitor. We show that foragers of the aggressive meliponine bee, Trigona spinipes, can detect and orient towards odour marks deposited by a competitor, Melipona rufiventris, and then rapidly take over the food source, driving away or killing their competitors. When searching for food sources at new locations that they are not already exploiting, T. spinipes foragers strongly prefer M. rufiventris odour marks to odour marks deposited by their own nest–mates, whereas they prefer nest–mate odour marks over M. rufiventris odour marks at a location already occupied by T. spinipes nest–mates. Melipona rufiventris foragers flee from T. spinipes odour marks. This olfactory eavesdropping may have played a role in the evolution of potentially cryptic communication mechanisms such as shortened odour trails, point–source only odour marking and functionally referential communication concealed at the nest.

A morphologically specialized soldier caste improves colony defense in a neotropical eusocial bee

Division of labor among workers is common in insect societies and is thought to be important in their ecological success. In most species, division of labor is based on age (temporal castes), but workers in some ants and termites show morphological specialization for particular tasks (physical castes). Large-headed soldier ants and termites are well-known examples of this specialization. However, until now there has been no equivalent example of physical worker subcastes in social bees or wasps. Here we provide evidence for a physical soldier subcaste in a bee. In the neotropical stingless bee Tetragonisca angustula, nest defense is performed by two groups of guards, one hovering near the nest entrance and the other standing on the wax entrance tube. We show that both types of guards are 30% heavier than foragers and of different shape; foragers have relatively larger heads, whereas guards have larger legs. Low variation within each subcaste results in negligible size overlap between guards and foragers, further indicating that they are distinct physical castes. In addition, workers that remove garbage from the nest are of intermediate size, suggesting that they might represent another unrecognized caste. Guards or soldiers are reared in low but sufficient numbers (1–2% of emerging workers), considering that <1% usually perform this task. When challenged by the obligate robber bee Lestrimelitta limao, an important natural enemy, larger workers were able to fight for longer before being defeated by the much larger robber. This discovery opens up opportunities for the comparative study of physical castes in social insects, including the question of why soldiers appear to be so much rarer in bees than in ants or termites.

Clustered male production by workers in the stingless bee Melipona subnitida Ducke (Apidae, Meliponinae)

Summary: In stingless bees brood cells are sequentially filled with liquid larval food (mass-provisioning), upon which the queen lays an egg. Thereafter the cell is closed by a worker. This study showed that during these processes workers of Melipona subnitida regularly laid eggs that served as food for the queen. Occasionally cells were oviposited in and immediately closed by a worker. These cells always rendered males. Some of these reproductive workers were seen to lay a trophic egg as well. Cells which were exclusively oviposited in by the physogastric queen gave rise to workers and queens only. In one colony it could be verified that three workers alone, which differed in age by one day, laid 15 male-producing eggs within a period of two successive weeks. Among them the number of ovipositions was positively related to the order in which workers eclosed - the oldest worker laying most eggs - and inversely related to the number of times they closed cells oviposited in exclusively by the queen. Apparently the physogastric queen was not able to stop certain workers from reproducing. We therefore conclude that some workers in M. subnitida temporarily dominated their queens in egg-laying.

Identification of oxygen containing volatiles in cephalic secretions of workers of Brazilian stingless bees

Os constituintes volateis das secrecoes cefalicas de 11 especies brasileiras de abelhas sem ferrao da linha Tetragonisca - Tetragona foram analisados. Foram identificados 145 compostos, por cromatografia gasosa e espectrometria de massas, dentre os quais 72 esteres, 22 alcoois, 16 acidos carboxilicos, 13 terpenos, 8 aldeidos, 4 compostos aromaticos, 2 lactonas e 1 diidropirano. As relacoes estruturais, origem e a distribuicao detes compostos sao discutidas. Com respeito a composicao qualitativa e quantitativa, cada especie mostra um buque de odores, os quais sao obtidos a partir de componentes poucos especificos. Em algumas especies proximamente relacionadas foram observadas similaridades no buque de odores. As fragmentacoes no espectro de massas de esteres graxos e dos esteres insaturados derivatizados com DMDS sao discutidas em detalhes.