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In 1977 ecologists Stephen Hubbell and Leslie Johnson recorded a dramatic example of how social interactions can produce and enforce regular spacing in a population. They studied competition and nest spacing in populations of stingless bees in tropical dry forests in Costa Rica. Though these bees do no sting, rival colonies of some species fight fiercely over potential nesting sites.

Stingless bees are abundant in tropical and subtropical environments, where they gather nectar and pollen from a wide variety of flowers. They generally nest in trees and live in colonies made up of hundreds to thousands of workers. Hubbell and Johnson observed that some species of stingless bees are highly aggressive to members of their species from other colonies, while other species are not. Aggressive species usually forage in groups and feed mainly on flowers that occur in high-density clumps. Nonaggressive species feed singly or in small groups and on more widely distributed flowers.

Hubbell and Johnson studied several species of stingless bees to determine whether there is a relationship between aggressiveness and patterns of colony distribution. They predicted that the colonies of aggressive species would show regular distributions, while those of nonaggressive species would show random or closely grouped (clumped) distribution. They concentrated their studies on a thirteen-hectare tract of tropical dry forest that contained numerous nests of nine species of stingless bees.

Though Hubbell and Johnson were interested in how bee behavior might affect colony distributions, they recognized that the availability of potential nest sites for colonies could also affect distributions. So as one of the first steps in their study, they mapped the distributions of trees suitable for nesting. They found that potential nest trees were distributed randomly through the study area. They also found that the number of potential nest sites was much greater than the number of bee colonies. What did these measurements show the researchers? The number of colonies in the study area was not limited by availability of suitable trees, and a clumped or regular distribution of colonies was not due to an underlying clumped or regular distribution of potential nest sites.

Hubbell and Johnson mapped the nests of five of the nine species of stingless bees accurately, and the nests of four of these species were distributed regularly. All four species with regular nest distributions were highly aggressive to bees from other colonies of their own species. The fifth species was not aggressive, and its nests were randomly distributed over the study area.

The researchers also studied the process by which the aggressive species establish new colonies. Their observations provide insights into the mechanisms that establish and maintain the regular nest distribution of these species. Aggressive species apparently mark prospective nest sites with pheromones, chemical substances secreted by some animals for communication with other members of their species. The pheromone secreted by these stingless bees attracts and aggregates members of their colony to the prospective nest site; however, it also attracts workers from other nests.

If workers from two different colonies arrive at the prospective nest at the same time, they may fight for possession. Fights may be escalated into protracted battles. The researchers observed battles over a nest tree that lasted for two weeks. Each dawn, fifteen to thirty workers from two competing colonies arrived at the contested nest site. The workers from the two colonies faced off in two swarms and displayed and fought with each other. In the displays, pairs of bees faced each other, slowly flew vertically to a height of about three meters, and then grappled each other to the ground. When the two bees hit the ground, they separated, faced off, and performed another aerial display. Bees did not appear to be injured in these fights, which were apparently ritualized. The two swarms abandoned the battle at about 8 or 9 A.M. each morning, only to re-form and begin again the next day just after dawn. While this contest over an unoccupied nest site produced no obvious mortality, fights over occupied nests sometimes kill over 1,000 bees in a single battle.

OFFICIAL32 The phrase “insights into” in the passage is closest in meaning to

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