C1139 Social Insects. The Society.

Lecture 4

The society: life cycle and kin structure

Aims
1. To provide information on colony life cycles and kin structure.

Objectives
1. To learn some of the diversity of colony life cycles in the Hymenoptera (e.g. how colonies are founded;
stages in the colony life cycle: founding, growth/ergonomic, reproductive, queenlessness/decline/death),
including specific examples.
2. To learn some of the diversity in the kin structure of colonies caused by variation in the number of queens
and the number of males to which queens are mated, including specific examples.

Big Picture
The previous lecture looked at the individuals within a society. The individuals are the building blocks of an
insect society. In this lecture we will consider the colony as a whole. A colony of eusocial insects is not some
random collection of conspecifics. Different taxa typically have well defined colony characteristics, such as the
way that a new colony is founded, the life cycle of the colony, the size it reaches, and the type of nest it builds.
We will focus on two key properties: colony life cycles and kin structure.

Colony founding by a single queen

Most multi-cellular organisms start life as a fertilized egg. Something analogous is also common in social
insects. The most common way for a hymenopteran colony to be founded is by a single inseminated queen.
This is sometimes called haplometrosis, but a more user-friendly term is single-queen founded. This is the
way that colonies are founded in most ants, Vespinae wasps, bumble bees (Bombus spp), and Halictidae sweat
bees. In termites, the most common method is by a mated pair. After a nuptial flight, termite kings and queens
rapidly pair up and look for somewhere to establish a nest.
In most ants, the newly-mated queen finds somewhere suitable to nest and snaps off her wings. She
rears the first batch of larvae using her body reserves (fat, flight muscles) which are converted into eggs or
oral secretions to feed the larvae. The first larvae often give rise to small workers known as minims or
nanitics. They then take over the work and forage. The queen herself does not forage. This is known as
claustral founding. (In some ants the queen does forage.) In bees and wasps the queen forages for food, and in
wasps for building materials as well.

Colony founding by multiple queens

This is called pleometrosis or multiple-queen founding. It can also be referred to as a "foundress
association" or "primary polygyny".
In many ants two or more unrelated ant queens found a nest together after a mating flight. Typically,
many queens make mating flights at the same time so that many queens are establishing nests at the same time,
and so can easily co-found. The queens co-exist until workers emerge, then they normally start to fight. As a
result, primary polygyny normally results in secondary monogyny. That is, the result is a nest with a
single queen even though it was founded by several queens. (In a few species the queens do not fight and co-
exist in the same colony.) Ant foundress associations are an example of mutualism. A foundress association
can help the nest get off to a rapid start, by rearing more workers, which helps the colony to survive the very
risky founding stage. In may ants there is much colony death during the founder stage due to intraspecific
competition and brood raiding. Many queens may establish nests in an area big enough for only a few mature
colonies. This is similar to what happens when many seeds sprout in an area too small for all the young plants
to survive.
To see how co-founding can be mutualistic, consider this example. A nest co-founded by two queens
has three times the chance of surviving the founding stage. Then the queens fight and one survives. Both
queens have an equal chance of winning. The average fitness of co-founding queens is then 1.5 times (3/2)
than of single-founding queens. The co-founding queens enter into a lottery in which the price of buying a
ticket is less than the average payout. If the co-founded nest had only 1.5 times the chance of surviving it
would not be worth co-founding.
In Halictidae bees and Polistes wasps, foundress associations often form in the spring when nests are
founded by overwintered females. In contrast to the ant situation above, the queens are often sisters who were
reared in the same nest and recognize each other (nestmate recognition) in the spring by their distinctive blend
of cuticular chemicals, some of which they acquired from their natal nest making them have a similar odour.
There is usually a dominant individual and one or more subordinates, who do more of the work and less of the
egg-laying.

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Colony fission: founding by queen(s) and workers

In many ants, one or more queens plus a group of workers establishes a new nest close to the existing nest. For
example, the clusters of wood ant (Formica lugubris) colonies that you may have seen on field trips to the
Surprise View at Hathersage, in the Peak District near Sheffield. This is typically associated with polygyny
(nests with multiple queens). You can sometimes see workers carrying brood walking along trails connecting
the nests.
In some species of ants, such as Eciton and Dorylus army ants, the queens have completely lost their
wings, and colonies are only formed by fission, never by a queen or queens alone.

Colony fission: founding by queen(s) and workers: swarming

In honey bees (Apis), stingless bees (Meliponinae), and Epiponini wasps (a tropical American group of
Polistinae wasps) colonies are founded by a swarm. That is, a mass of workers who leave the parental nest
with one or many (Epiponi) queens. Swarming has evolved multiple times. In many swarming and budding
species the colony cannot function without workers. A honey bee colony cannot function at all with less than
approximately 1000 workers. An Ecition army ant colony cannot function with less than approximately
100,000 workers. In this species, their social life has evolved to such an extent that small colonies cannot
survive.
Swarming can also be used for colony relocation, without colony fission. The whole colony leaves an
unsuitable nest site. African honey bee, Apis mellifera, and Asian giant honey bee, Apis dorsata, swarms may
even be migratory, with the whole colony travelling up to hundreds of kilometers to a more favourable location
in terms of flowers. In Apis dorasata the migration is seasonal with the swarms returning several months later
to where they originally set out from. Amazingly, an Apis dorsata colony nesting in one particular tree may
leave, and come back to exactly the same tree months later.

Reproductive swarming in the honey bees and stingless bees

This usually occurs in spring in temperate areas, late April to June in England. The mother queen and
approximately two thirds of the adult workers leave the nest in the "prime swarm". It takes just a few minutes
to leave the nest. The bees form a cloud in the air, but rapidly settle in a cluster usually on a nearby bush. (At
the time the swarm departs the original nest contains c. 10-20 pupal queens. When these hatch none, one, or a
few additional swarms, known as casts or after swarms, may depart each headed by a virgin queen. One
of the newly-reared queens takes over the original nest.) Scout bees on the swarm now look for cavities in trees
and other locations as potential nest sites. One is chosen, usually within 1km, by a complex decision making
process in which a whole range of parameters, including cavity volume, height above ground, size of cavity
entrance, and entrance orientation are evaluated. The swarm then moves to the new cavity and in just a few
minutes the bees, on average about 10,000, all pile in. Overnight, using wax from their wax glands, the colony
will build its first comb and the queen will start laying eggs. The swarm-founded colony is in a race against
time to build up its population and honey stores before winter. In northern areas most swarm-founded colonies
die out in their first winter due to starvation.
In stingless bees (Meliponinae) swarming is more gradual. A colony locates a suitable nest site and
gradually builds combs and food pots while maintaining contact between the new nest and the old nest.
Eventually, a young queen, not the mother, flies over to the new nest to join the workers there. The process can
take weeks, even months. Stingless bees and honey bees are close relatives. But the large differences in their
swarming behaviour suggest that swarming evolved independently in each group.

Parasitic nest founding

It is common for queen Vespinae wasps and bumble bees to steal a nest being founded by another queen of the
same species. In addition, both Vespinae wasps and bumble bees have species that are workerless parasites and
take over the nest of other species. The workers of the deposed queen rear the young of the usurping queen
even though they are of another species. In Britain, the cuckoo wasp Vespula austriaca is a workerless
parasite of Vespula rufa, the red wasp, and there are also several parasitic species of bumblebees in Britain.
There are also workerless ants parasitic ants. In addition, in many ants that do have workers a queen may
found her nest by taking over the nest of another species.
In the workerless parasitic species eusociality has gone over to parasitism. It should not be a surprise
that social parasitism occurs. Workers can rear the offspring of another female, in just the same way that
parents can, as in the cuckoo. In some ants, such as the British wood ant Formica sanguinea, workers raid
colonies of other wood ant species and capture pupae. The pupae emerge in the F. sanguinea nest and work
even though they are helping to rear brood of another species. This is known as slavery or dulosis. Workers
in some ant species, such as Polyergus, cannot work. All they can do is raid other nests. All the work is done

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by the allospecific slaves. In F. sanguinea workers can still work if they have to. Anyone interested can read in
Origin of Species some simple experiments that Charles Darwin did on F. sanguinea.

Colony life span: Annual

Annual nests are founded in spring and die out in the summer or autumn. The colony first rears workers then
queens and males. This is a common life cycle and occurs in Bombus, Vespinae wasps, Polistes wasps, and
most eusocial Halictidae bees. During winter all that remain are young inseminated queens. The young queens
and males are reared in the summer or autumn, leave the parental nest, and mate. The males die and the
inseminated females overwinter.

Colony life span: Perennial

Perennial means a colony that can live for more than one year. Colonies of termites, ants, Apis and
Meliponinae bees, some Halictidae bees and Epiponini wasps are perennial. In these bees and wasps and some
of the ants the colony can replace the queen if she dies. These colonies are potentially immortal, but still perish
by being predated, starving, or by something going wrong during the queen replacement process.

Four stages of the colony life cycle

Oster and Wilson (1978) characterize four stages in the life of an ant or social insect colony. Again, there are
analogies to the life cycle of a multi-cellular organism.
Founder Queen(s) and brood only.
Ergonomic Workers present. Colony is growing but rearing only workers. Lasts just a few months in
annual species, but can last several years in large-colony ants such as Atta leafcutter ants. In perennial
colonies there may be an ergonomic phase and a reproductive phase each year.
Reproductive Colony is producing queens and males.
Orphaned/queenless A colony in which the queen has died and cannot/has not been replaced. In many
species (e.g. Apis mellifera) the workers now lay many eggs that are reared into males. Because no female
eggs are being laid the workforce dwindles and the colony dies out. In some annual species (Bombus, Vespa,
Dolichovespula) workers may actually kill the queen and take over egg laying.

Colony kin structure

We will return to this in a later lecture in the second section of the lecture course. Today we will just touch on
this important topic. Kin structure can be a description of a colony in terms of family relationships. For
example, the workers are all offspring of a single queen mated to a single male. It can also be a description in
terms of relatedness among specified individuals. For example, the relatedness among workers is 0.75. Note
that different kin structures may give the same relatedness. For example, both multiple mating by a single
queen and multiple queens reduce relatedness among workers.

Examples
The simplest colony kin structure is a single queen mated to a single male. All workers are the offspring of this
pair. This is a very common type of kin structure, found in many ants (e.g. the fire ant Solenopsis invicta),
almost all Meliponinae bees, bumble bees, and Polistes wasps. In some of these species the males are all the
queens sons, and in others some or many young males are also workers sons.
It is also very common to have a single queen mated to more than one male so that the workers all
have the same mother but have two or more fathers. In many species most queens are mated to one male, with
a small proportion being mated to two males (e.g., in some Vespinae wasps such as the hornet Vespa crabro,
Dolichovespula). In common wasps, Vespula vulgaris, the queen is normally mated to several males.
Extreme multiple mating by queens (>5 males) has evolved multiple times and is is found in a few
taxa including Apis honey bees, Dorylus and Eciton army ants, Acromyrmex leafcutter ants, Pogonomyrmex
seed harvester ants, Cataglyphis desert ants, and some Vespula wasps. In Apis the workers have one mother
but 10, 20 or even 50 fathers.
Why is there so much variation in queen mating frequency and why has multiple mating evlved on
some species? A lot of work has been done on this topic but we will not cover this topic in the course.
In many ants, colonies have more than one egg laying queen. This is also common in Epiponini wasps
but not common in bees or other eusocial wasps.