Abril, 1997 An. Soc. Entomol. Brasil 26(1) 153 Labor Division in Melipona compressipes fasciculata Smith (Hymenoptera: Apidae: Meliponinae) Katia M. Giannini1 1Departamento de Biologia, F. F. C. L. R. R, USP, Av. Bandeirantes, 3900, 14040-901, Ribeirão Preto, SP. An. Soc. Entomol. Brasil 26(1): 153-162 (1997) Divisão de Trabalho em Melipona compressipes fasciculata Smith (Hymenoptera: Apidae: Meliponinae) RESUMO - A divisão de trabalho em Melipona compressipes fasciculata Smith foi analisada e comparada com outras abelhas eusociais - Meliponini e Trigonini. Os resultados mostraram que existe uma sequência de atividades similar a outros meliponíneos já estudados, a saber: “grooming”, trabalho com cerume, construção de células, limpeza da colônia, participação no processo de apro- visionamento e oviposição (POP), desidratação de néctar, ventilação, vedação de frestas, recepção de néctar, guarda e forrageamento. PALAVRAS-CHAVE: Insecta, Meliponini, abelhas sem ferrão, divisão de tra- balho, abelhas eusociais. ABSTRACT - Labor division in the stingless bee Melipona compressipes fas- ciculata Smith was analyzed and compared with that of other eusocial bees - Meliponini and Trigonini. The results showed a sequence of activities which was similar to that of other stingless bees already studied, such as: grooming; cerumen work; cell construction; cleaning; the colony; provisioning and ovi- position process; ventilation; enclosure gap; nectar reception; and foraging. KEY WORDS: Insecta, Meliponini, stingless bees, division of labor, eusocial bees. The existence of physical and temporal cas- apud Winston & Fergusson 1986, Donhoff tes is one of the most complex and important 1855 apud Winston & Fergusson 1986). Fur- factors in the social structure of insect colonies ther research analyzed the flexibility in tempo- (Oster & Wilson 1978). The physical castes in- ral labor division and the underlying glandu- clude a queen, a small number of males, and the lar basis for tasks performance (Rösch 1925 numerically predominant worker caste. The tem- apud Winston & Fergusson 1986, Ribbands poral caste consists of temporal labor division 1952, Lindauer 1953, Sekiguchi & Sakagami in which there is an ontogenetic sequence of 1966). Recent investigation has involved the activities performed during the workers’ lifetime. colony-level factors which try to determine In the earliest studies on labor division of the reasons why bees perform duties at diffe- honeybees, it was reported that worker acti- rent times (Free 1961, Kolmes 1985,Winston vities presented a temporal basis (Butler 1609 & Fergusson 1985, among others). 154 Giannini The typical labor division in eusocial bees were measured based on the number of brood proceeds in the following sequence: callow, combs and on the number of pollen and honey nursing, household, and foraging (Sakagami pots. The 1st observation was carried out in 1982). However, there is considerable flexibi- the beginning of the experiment and the 2nd lity concerning the age-related trend of acti- when the experiment was finished (Table 1). vities so that the tasks sequence can be adjus- Observation was carried out in both extra ted in relation to the colony external or internal and oviposition periods - i.e., 8h of daily requirements. observation. The marked workers were recor- Some of the research on labor division in ded as performing one or more tasks through Meliponinae shows the sequence of tasks individual observations. performed during the workers’ lifetime (Bas- sindale 1955, Kerr & Santos Neto 1956, He- bling et al. 1964, Darchen 1969). More recently, Table 1. General conditions of Melipona research has described a temporal sequence compressipes fasciculata colonies. of labor division in Melipona bicolor bicolor Lepeletier (Bego 1983), Melipona favosa Fa- bricius (Sommeijer 1984), Frieseomelitta languida Moure & Camargo (Ribeiro 1989), Nannotrigona (Scaptotrigona) postica La- treille (Simões & Bego 1991), Tetragonisca angustula angustula Latreille (Grosso 1993) and Plebeia remota Holmberg (van Benthem et al. 1995), among others. This is a contribution to add more infor- mation on the activity of individual workers in relation to the temporal tasks sequence of Melipona compressipes fasciculata Smith, comparing the results with other Meliponinae species. Materiais and Methods This study was conducted at Ribeirão Preto, S.R, Brazil, from September to December 1991. Two colonies of M. compressipes fasci- culata from São Luís, MA, were used. Colo- The activities performed by workers as nies were manipulated only for the introduc- well as queen/worker interactions were obser- tion of marked workers. A marked cohort of ved according to the following classification: 111 newly emerged workers was introduced Grooming = a bee cleans any parts of its body into an observation hive. The cohort was ob- with its mouthparts or legs; Cerumen work = tained by removing combs from another hive removal of cerumen from old brood cells, cons- so that bees could emerge overnight in an in- truction of support pillars, food pots and invo- cubator at 28°C to be later introduced into the lucrum; Cell construction = mainly mandibular hive 1-2h after marking. During 14consecutive ”manipulation” of cell parts or working inside days all the workers which emerged were in- the cell; Provisioning and oviposition process dividually marked. These workers were up to (POP) = participation in the provisioning and 24h old. The general conditions of the colony oviposition process of brood cells (Arousal Abril, 1997 An. Soc. Entomol. Brasil 26(1) 155 stage, food discharge, worker oviposition and Convex curves are found in species with low cell operculation); Cleaning = manipulation mortality during the initial phases of life. The of waste material and pieces of cocoon that high mortality was observed to be true of age are carried out of the colony; Enclosure gap brackets in which workers became foragers. = with cerumen and mud; Ventilation = wing In the other species of Meliponinae bees beatings; Trophailaxis (Tr) = food exchange the following longevities were recorded: Ple- from worker to queen; Food reception = from beia (Friesella) schrottkyi Friese, 30.1 days worker to worker; Beg food (B) = from queen (Camillo-Atique 1977), F. languida, 33.3 days to worker; Nectar reception = from worker to (Ribeiro 1989), M. bicolor bicolor, 44.0 days worker; Foraging = collection of food and ma- (Bego 1983), N. (Scaptotrigona) postica for terial for nest construction; Insertion (I) = fo- three colonies estudied, 26.3 days, 39.5 days and 32.9 days (Simões & Bego 1991), M.fa- rebody insertion into cell; Escape (E) = worker vosa, 40.0 days (Sommeijer 1984) and Me- escapes when the queen approaches; Advan- lipona scutellaris Latreille, 27.4 and 34.5 days ce (AD) = simple advance (a worker darts on (H. H. Oliveira & A. Kleinert-Giovannini, pers. the queen); Advance-back (AB) = the worker inf.) for two different colonies observed. The performs advance and back movements in internal conditions of the colony, observations the presence of the queen. in different periods of the year, and different morphological and physiological conditions Results and Discussion among the species are probably responsible for these variations. The average longevity observed for M. When analyzing our data according to compressipes fasciculata was one of 42.5 (Ito the workers’ age sequence, the main aspects 80 days) (Fig. 1). According to Sakagami & described below were observed: the cerumen Fukuda (1968), the type of survival curve is work is one of the 1st activities performed by directly correlated with the division of labor. workers in M. compressipes fasciculata.The Figure 1. Survival curve (lx) and mortality curve (dx) for workers of Melipona compressipes fasciculata. 156 Giannini Figure 2. Frequency of workers of Melipona compressipes fasciculata engaged in vari- ous activities according to respective age in brackets. Abril, 1997 An. Soc. Entomol. Brasil 26(1) 157 grooming task and colony cleaning are activi- 1983). Thus, the internal conditions related to ties that overlap almost completely. Duties as the number of workers available to perform reception, nectar dehydration and foraging each task, the queen’s laying rate, stored food, are performed only after the bees have partici- the age of colony members, etc. are important pated in the cell construction and oviposition factors in stablishing such dynamic. External process. Trophailaxis and food reception conditions such as seasonality (Ceccato 1970, occur ali during the workers’ lives. It is impor- H. H. Oliveira & A. Kleinert-Giovannini pers. tant to consider that trophallaxis is more cons- inf), availability of food resources and nest picuous when bees are older, and that nectar construction material, etc. can also influence reception starts when they are young. Beg colony dynamic. food is performed throughout the bees’ lifetime A important point is the overlap of duties. (Fig.2). A determined worker can perform more than The activities observed by different au- one activity at the same time in all the species thors do not agree with those defined for M. observed. The frequency of bees’ performing compressipes fasciculata, which makes the different activities throughout their lives (Fig. comparison of results difficult. It is important 3) were: 99.1% worked with cerumen; 76.6% to emphasize the fact that labor division is worked in cell construction; 52.3% partici- dynamic and that it varies with species, colo- pated in the oviposition