7 How stingless bees solve traffic problems Jacobus C. Biesmeijer E. Judith Slaa Dirk Koedam KEY WORDS Melipona, Trigona, traffic flow, flight activity, social insect, nest architecture Entomologische Berichten 67 (1-2): 7-13 In this paper we examine to what extend the size and shape of stingless bee nest entrances can be explained as being a compromise between traffic flow optimization and adequate defence. Small bee colonies have small entrances that are easily defendable. The trade-off is that traffic jams and collisions are common at very high foraging activity. The shape of the entrance of large bee colonies tends to be such that there is limited overlap in take-off and landing areas and, as a result, no or few traffic jams. Large entrances are probably easily located by predators and parasites, which may explain why such colonies are highly defensive. Introduction the entrance plays a role in traffic flow. Throughout the paper The majority of human city-dwellers experience daily problems we compare the architectural solutions used by stingless bees in traffic. Cars congest the roads leading into and out of urban with human solutions for similar problems. centres and pedestrians queue at stations, elevators and super- market checkouts. Some of these problems arise accidentally, Stingless bee cities: a short introduction for example as a result of parked cars or road works, but many problems occur at human-designed infrastructure, for example Stingless bees live in colonies of several hundreds to tens of one-lane bridges, car parks, crosswalks and building entrances. thousands of workers headed usually by one reproducing This seems to suggest that, although engineers and architects queen. Since all workers are offspring of the queen and most do base the planning and design of human infrastructure on ample not participate in reproduction (Koedam et al. 1996, 2001), their study of car movement and behaviour of pedestrian crowds, interests concerning foraging, colony growth and maintenance they are not able to keep up with the changing demands of our are largely aligned. Therefore, we might expect that the design daily life and traffic problems remain. To most of us, waiting in of the nest reflects the requirements of the colony as a whole line seems an unavoidable part of life, but it might be a fairly re- and not the individual preference of one or few architects (note cent phenomenon; traffic jams were probably far and between the difference with human cities). In addition, the nest building in cavemen societies and in times when most transportation and maintenance is a continuous and self-organized process. was by horse-drawn carriage. This means that no leader is giving orders (unlike in human Modern humans in densely-populated areas are not the only building activities) and that no blueprint or recipe is followed ones to experience congestion problems. Long before early hu- strictly (unlike some burrow excavation by solitary insects or mans left Africa, other group-living animals, the social insects, web building by spiders), but that hundreds of individual worker had to live with high-volume traffic flow. Among the oldest fos- bees make their own decisions on where and how to change the silized social insects are members of now extinct taxa of the nest structure. These individual decisions are based solely on tropical stingless bees (Hymenoptera, Apidae, Meliponini). The local information obtained by builders that lack a big plan (and remarkable thing is that these amber bees are workers, sterile possibly some genetic predispositions) [see papers in Detrain et females, which were trapped in tree resin probably while col- al. (1999) for an introduction on the role of self-organization in lecting this material for nest construction. These workers were social insects]. citizens of insect cities with division of labour between an egg- Since foraging takes place outside the insect city, entrances laying queen and the workers that built and maintained the of sufficient size and number are needed to facilitate traffic nest. Densely-populated insect societies have been around for flow. The larger and more conspicuous the entrance, the easier at least 80 million years (e.g. Engel 2001). the bees move in and out of the colony. On the other hand, pre- Like human cities, social insect cities have complex internal dators and parasites benefit from large entrances, and pressure infrastructure and, sometimes, external trail systems (in ants to limit their access may have driven the size of the city’s en- and termites). The link between the city environment and the trances down. Similarly, maintenance of high temperature and outside world are the city’s entrances. They facilitate the flow of not too low humidity in the nest is essential for optimal brood incoming and outgoing traffic. In this paper we investigate how development. We can expect that nest entrance architecture, one group of social insects, the stingless bees, organizes traffic e.g. size, number, shape and conspicuousness, has been under flow in and out of their colony. First, we analyze whether the pressure of natural selection and that the design is a compromi- size of the nest entrance is correlated with colony size. Second, se between foraging requirements, adequate defence and main- we test whether species-specific entrance size results in ade- tenance of internal conditions. If this is the case, the entrance quate traffic flow. Third, we investigate whether the shape of might represent a bottleneck to the colony’s traffic flow. We test 8 entomologische berichten 67(1-2) 2007 a b attack the intruder (level 4). We correlated defensiveness with relative entrance size and found that all bees with large entran- ces (>10 bee heads wide) are defensive (at least level 2), whereas bees with small entrances (<10 bee heads wide) can display any level of defensiveness, from being timid to being easily excited by disturbance (figure 3). Traffic jams and collisions c d The observed correlations seem to suggest that the entrance size may indeed be a compromise between adequate defence and smooth traffic flow. If this is the case, then having a small entrance should be costly and traffic flow should be hampered, especially if foraging activity is very high. Traffic problems might include traffic jams or collisions. To assess whether traf- fic jams are more prominent at high traffic flow rates we video- Figure 1. Some examples of stingless bee nest entrances. a The recorded arriving and departing bees at the nest entrance of 16 one-bee-wide entrance of a Melipona quadrifasciata Lepeletier colony with a spider lurking nearby, b four bees fit in the entrance of a stingless bee species at various traffic flow rates. Undisturbed Scaura latitarsis (Friese) nest, c a funnel entrance with a large lower traffic is characterized by a random sequence of bees entering lip of a Tetragona perangulata Cockerell nest, d Scaptotrigona pectoralis and leaving the nest, whereas a non-random ‘aggregated’ se- tend to have a tubular opening with a small lower lip. Photos: Koos quence of bees going in and out is indicative of traffic jams. Biesmeijer Enkele voorbeelden van nestopeningen van angelloze-bijennesten. a Slechts Think of how we pass road works that leave only one lane open. één bij tegelijk past in de opening van dit Melipona quadrifasciata-nest, b Several cars will pass from one direction, followed by a series of vier bijen tegelijk passen in de opening van dit Scaura latitarsis-nest, c de cars from the other direction and so forth. When arriving at trompetopening met grote onderlip van een Tetragona perangulata-nest, such a point we try to take advantage of the flow by quickly fol- d Scaptotrigona pectoralis heeft een buisvormige opening met een zeer kleine onderlip. lowing the car in front of us passing the blockade and making passing impossible for the cars at the opposite end. In stingless bees with narrow nest entrances a similar situation might deve- this hypothesis with tropical stingless bees. The several hund- lop at high traffic flow rates. Pressure of bees moving in and out red species present a stunning array of species-specific nest en- builds up and the side with highest pressure (i.e. most bees pu- trance designs (Biesmeijer & Slaa 2003; see also Wille & Miche- shing forward) wins and those bees move. This lowers the pres- ner 1973, Michener 1974, Camargo & Pedro 2003), the function of sure and allows the other side to move so that a series of in- which has not been explained (figure 1 shows some examples). and outgoing bees alternate. We analyzed the sequence of arri- vals and departures using the runs-test (Siegel & Castellan 1988). Explaining variation in entrance size We can derive two predictions from our hypothesis that en- entrance capacity (n bee heads) trance size is a compromise between colony size (to facilitate traffic flow), defence (to limit predation and parasitism) and nest microclimate. The first prediction is that colony size and total entrance size are positively related. If there are no costs to a large entrance size, small colonies could have large entrances without having problems defending the colony, and large colo- nies could have small entrances without having traffic flow pro- blems. A review of published records and personal observations revealed that entrance size increases with colony size (figure 2). Colony size explains 52% of the variation in entrance size. Addi- tional support for the hypothesis that the entrance is shaped by traffic and defence, is that many of the species that are posi- tioned high above the regression line are very defensive (figure 2, abbreviations Om, Po, Tc) and thus can afford a larger en- trance size. On the other hand, many of the species positioned colony population size (n individuals) below the regression line (e.g.