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The Polygyny of Melipona Bicolor: Scramble Competition Among Queens

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The Polygyny of Melipona Bicolor: Scramble Competition Among Queens Apidologie 37 (2006) 222–239 222 c INRA/DIB-AGIB/ EDP Sciences, 2006 DOI: 10.1051/apido:2006025 Original article The polygyny of Melipona bicolor: scramble competition among queens Hayo H.W. Va,HanD Vb,VeraL.I–Fc a Klemit 1, 5325 KG Wellseind, The Netherlands b Department of Behavioural Biology, Utrecht University, The Netherlands c Departamento de Ecologia, Instituto de Biologia, Universidade de São Paulo, Brazil Received 23 December 2005 – Revised 14 February 2006 – Accepted 15 February 2006 Abstract – The stingless bee Melipona bicolor is facultatively polygynous, a unique character among the bees. Polygynous colonies were not more productive than monogynous colonies. During the process of provisioning and oviposition of cells (POP) a queen may be either alone or together with one or two other queens. If together, each queen has on average the same chance to lay the egg, indicating that there is no dominance mechanism involved. During the POP, a queen may ingest some of the larval food and a trophic egg laid by a worker. Worker egg laying is less frequent in multiple queen POPs. The most active queen has proportionally more single-queen POPs and more trophic eggs. Such nutritional advantage and the resulting output of eggs could depend on chance, but a lasting qualitative difference among queens probably exists as well. Though we could outline the mechanisms behind the outcome of this scramble competition for egg laying, the adaptive significance of this polygyny remains largely mysterious. polygyny / stingless bee / Melipona bicolor / scramble competition / trophallaxis / trophic eggs / Apidae / Meliponini 1. INTRODUCTION a transient situation. It often has several laying queens in the colony, living together for con- Nearly all species of social bees are char- siderable periods of time. acterised by colonies that are headed by a Polygyny is not uncommon in ants, and single queen. Polygyny, the presence of sev- has been studied relatively well in a num- eral laying queens, whenever it occurs in ber of species. Two different ontogenies ex- bees, is only temporary. A well-known ex- ist in the ants. The first form of polygyny is ample is the replacement of an old or dam- cooperation among queens upon founding a aged queen in the honeybee colony by a young colony (pleometrosis), a cooperation that en- one. The two queens may live together for a hances the success in the early period. When limited period of time, sometimes both lay- the colony grows, its polygyny develops into ing eggs. Similar transient conditions have a functionally monogynous situation, either been found in some species of stingless bees with a dominant queen that lays the eggs and a (Apidae, Meliponini) (da Silva, 1972; Witter queue of other inseminated queens on a wait- and Wittmann, 1997). Melipona bicolor,a ing list or in a truly monogynous condition. stingless bee species from Southern Brazil, In the second form there is secondary poly- however, is a facultative polygynous species, gyny, where monogynous colonies accept in- where the cohabitation of laying queens is not vading new queens and where subsequently more than one queen is involved in the pro- Corresponding author: H.H.W. Velthuis, duction of sexuals. In both situations the de- [email protected] gree of relatedness among the queens plays a Article published by EDP Sciences and available at http://www.edpsciences.org/apido or http://dx.doi.org/10.1051/apido:2006025 Polygyny of Melipona bicolor 223 role in our understanding of the evolutionary of the species, colony founding, which is by origin of polygyny. swarming, might be difficult due to a shortage Hölldobler and Wilson (1990) and Keller of vacant nesting sites. Consequently, in many (1993) reviewed the adaptive significance of species swarming is rather infrequent. This is and the ecological conditions that favour particularly true for most Melipona, a neotrop- polygyny in these ants. In a fragmented habi- ical genus that contains about 40 species, of tat, where the population of an ant species is which 35 are found in Brazil (Silveira et al., subdivided into small and isolated local pop- 2002). It is only in M. bicolor that we find ulations, such mini-populations would bene- polygyny. What is it that distinguishes M. fit from the increased effective population size bicolor from its congeneric relatives? What (increased genetic variation) that goes along conditions could explain the occurrence of with polygyny. In contrast, if genetic variabil- polygyny exclusively in this species and why ity in small isolated populations was small, do polygynous and monogynous colonies co- altruistic cooperation would be favoured. In- occur? Before addressing questions like these, deed, the intuitive assumption that the queens we should definitely know more about the ba- in polygynous colonies are related often holds sic biology of the species and about prox- but is not universally supported. Apart from imate mechanisms that regulate social pro- selection at the levels of direct and indirect fit- cesses within the colony. ness of the individuals, colony-level selection All Melipona species have well-protected and population-level selection appear to be in- nests inside a pre-existing cavity, which can be volved. entered only through a long, narrow and strong Among the ecological conditions that may entrance tube penetrating deep into the nest. favour polygyny in ants we find: (a) the diffi- In the nest cavity itself two parts can be dis- culty in founding a colony for some species tinguished, a part containing the several lay- because of nest-site limitation and predation ers of horizontally arranged combs enclosed in on gynes, a factor that would favour the se- a well-developed involucrum and, outside the lection for polygyny in Formica (Rosengren involucrum, a number of food pots in which et al., 1993); (b) the abundance of food (in large amounts of food may be stored in sep- Myrmica, Herbers, 1993), where territorial de- arate honey and pollen pots. Because of the fence breaks down under abundant food sup- large amounts of food they produce, a num- ply, with the consequence that colonies be- ber of these species are kept in managed hives, come polygynous and the relatedness of work- a factor that in turn has promoted the study of ers with the sexuals that are produced in their their biology. colony drops. However, as Herbers (1993) al- The brood combs of Melipona are con- ready concluded, the role of ecological fac- structed sequentially and are removed once tors as the prime cause for the emergence and the brood has emerged from the cells. New maintenance of polygyny in ants is still poorly cells are added at the margin of the youngest understood. comb(s). Their construction, and especially Would this information on polygyny among the mass provisioning, egg laying and cell clo- the ants help us understand the polygyny in M. sure is a process involving a number of indi- bicolor? So far, the answer seems to be neg- viduals that engage in intense social interac- ative. The stingless bees form a diverse group tions. This process, known as POP (Provision of several hundred mainly tropical species, be- and Oviposition Process), has been described longing to about 25 genera (Michener, 2000). for many species (for reviews, see Michener, The nests are in pre-existing cavities such as 1974; Sakagami, 1982; Zucchi, 1993; Zucchi in the trunks and main branches of trees, in et al., 1999). abandoned arboreal termite nests, or deep un- Two subspecies of M. bicolor, M. b. bi- derground in abandoned ant and rodent nests. color Lepeletier and M. b. nigra Lepeletier In most species, the colonies are long-lived, have been distinguished. For the latter the syn- persisting perhaps for decades, while individ- onym M. b. schencki Gribodo exists. They dif- ual queens may live for 1–3 years. In many fer in coloration and in the geographic area of 224 H.H.W. Velthuis et al. their distribution. In the older literature these we describe these behavioural aspects in a qualita- three names were used for separate species. M. tive sense. Quantitative data were collected on the bicolor is a bee from forested areas, where it participation of each of the queens of a polygynous nests in tree cavities near the soil, usually at colony in these POPs, the frequencies with which damp places near a river or rivulet (Nogueira- they were involved in egg laying and their consump- Neto, 1997). The occurrence of polygyny in tion of the different types of food. this species was first noted by Kerr (1949), but The observations were made during the months this aspect has not been previously studied. In March–April and October–November. Altogether, fact, only a few studies have been published on five polygynous colonies were observed intensively. this species. Bego (1983) reported on several During these observations the cover glass of the bionomic aspects and da Silva et al. (1972) ex- outer box had to be removed to enable a better view; perimented with artificial insemination of its as a consequence, the temperature control was in- queens. According to the latter authors, the terrupted. Fortunately, in M. bicolor new combs are species can be kept in the laboratory without always constructed on top of the pile of already ex- isting combs; when at the bottom of the pile the great difficulty, provided the colonies succeed bees have emerged from the lowest comb, the en- in maintaining good control of humidity in the tire pile slowly sinks. Therefore, in this species it is nest. unnecessary to dismember the pile of combs to ob- In this paper we investigate the cohabita- tain access to the cell construction region. To get a tion of queens.
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