The Song of the Queen

The mating flight of a honey bee queen is orchestrated by events that occur days before she actually flies from her nest in pursuit of a gathering of drones, 20 or so of which will get lucky and mate with her. Her mating song begins within the nest while she’s still held captive in her royal cell by her worker sisters, piping and tooting audible sounds to her rival queens and her captors. The song changes pitch and intensity after she’s released from captivity and seeks out and destroys her rivals. The orchestra is joined by the workers running through the nest buzzing their wings, exciting the colony and the new virgin, pushing her toward the entrance, where she takes flight. When she finds the congregation of drones, they join in with the buzzing of their wings and popping sounds of their genitalia as they find and mate with her. The queen returns to the nest, the heir of the nest and colony; and a new superorganism is born.

How Bees “Paint” Their Local Environments

Bees are able to exploit their environments temporally as well as geographically. The results are like brushstrokes on the landscape. The effects of foraging activities of honey bees can be seen by the distribution of flowers. Honey bees have evolved mechanisms to forage optimally both as individuals and as colonies, expending their energy and short lives to maximally provision their nest. Their communication system directs foragers to different floral patches at different times of day, reflected in the changing kaleidoscope of color on the legs of returning pollen foragers. Their pollination transforms the landscape in the spring into splashes of color resembling a painter’s pallet.

The Superorganism

Insect societies have been likened to superorganisms since the early 20th century because they are organized around defense, nutrition, and reproduction, like our own bodies. Like individual organisms, they undergo development and separate the germ line (eggs and sperm in our case) from the body cells, the soma. In social insects, the germ line is sequestered in the reproductive individuals, while the body cells are the non-reproductive workers. The superorganism was proposed by William Morton Wheeler as a real entity but instead was used primarily as a metaphor, a structure for hanging analogies with human organisms and societies. Throughout the 20th century there were many twists and turns in the definition and use of the superorganism concept and many questions regarding its usefulness.

Environmental Artists

One hundred twenty-five million years ago, the earth exploded with color with the rapid evolution of flowering plants. The explosion coincided with the rapid increase of species of bees. The bees and flowering plants were locked in a dialectical dance of coevolution, each becoming adapted to the other. The flowers evolved to exploit the feeding habits of bees, and bees evolved to rob the flowers of their precious loads of pollen and nectar. Bees became social and developed communication and navigational systems to better exploit their environment. They continue to transform our world through their effects on the agricultural landscapes and the food we eat. But today the honey bee is threatened. Populations are declining, a consequence of commercial beekeeping and agricultural practices.

How a Superorganism Evolves

Superorganisms are reverse-engineered by natural selection. The totality of the organizational structure of a colony, its colony-level phenotype, is exposed to natural selection. Those colonies with phenotypes that are best adapted to their environment survive best and reproduce the most reproductive offspring. The heritable features of those colony phenotypes increase in frequency in the next generation, and the population evolves a social organization. Natural selection doesn’t “see,” doesn’t act on, the individual components of organization, only the gross product. Therefore, the intricacies of the design, such as reduced fertility of workers, nest design and maintenance, and defensive behavior, are reverse-engineered. They evolve as a consequence of their effects on the whole colony.

How to Make a Superorganism

Insect superorganisms are characterized by a reproductive division of labor (drones, queens, and workers) and a complex division of labor among the non-reproductive individuals, the workers. In the social bees that have attained the highest degrees of sociality, at or approaching superorganism status, males don’t survive mating and are only present as reproductive sperm sequestered in the queen. Queens and workers are anatomically differentiated but derived from the same genome. Differentiation is a consequence of differential feeding of developing larvae by the workers. In the honey bee, worker nurse bees manipulate the developing larvae, forcing them into their reproductive roles. The adult workers self-organize into an ordered society, performing all of the functions necessary for colony survival and reproduction. There are no task masters or forewomen directing the workforce. Instead, every individual makes local decisions about their behavior based on their response thresholds to stimuli in their environment.

Reproductive Competition

Academic requirements for an insect society to be considered a superorganism vary broadly with authors and among sociobiologists. The two conditions that emerge as the most important are the lack of reproductive competition among nestmates and the state of social evolution where workers within colonies are at or beyond the “point of no return”—workers have lost the ability to live and reproduce independently. The search for reproductive competition within highly social colonies of ants and bees is a large academic research enterprise filling the contents of countless journals and books. However, a look at honey bees shows that these special conditions not only aren’t met but obfuscate the wonderful array of social states in which colonies exist within one population.

The Social Contract

Animals that live together in a society, like social insects, have a tacit agreement, a social contract of sorts, that guarantees that their reproductive interests are protected in exchange for their social cooperation. This contract isn’t written on paper, nor is it expressed in explicit laws or national constitutions; but instead it is written into the DNA of populations with the ink and quill of inclusive fitness and natural selection. All social groups share common features of providing for the defense of social members from external threats, internal policing of cheating by those who don’t cooperate, and some kind of protection of reproductive rights, either direct or indirect. Social benefits of insect societies include organizational structures similar to those of human societies such as public works, public health, police, and border patrol. Without these features, they would fail as societies.

Environmental Engineering

Bees engineer the environment. Their foraging activities change the floral composition near the nest, thereby changing the niches of other species that depend on the vegetation for food and shelter. Changes in floral abundance and composition resulting from their activities may also benefit them directly or descendant colonies. Honey bees also engineer their own environment by constructing a protective nest. The nest of the honey bee provides protection from the external environment by providing an insulated shell within which they live and wax comb to serve as a substrate for social interaction, food storage, and a nursery for raising larvae. They have community systems for healthcare, thermal regulation, and defense.