The social structure of insect societies

What is different between insect

societies and swarms of locusts,
fish or birds?
 The social insects have royalty!
• Social Hymenoptera (ants, wasps, bees) have a queen
• Social Isoptera (termites) have a king & queen
• The queens (and kings) are typically long lived (15 years
in termites, 4 years in honeybees)
• Their sole task is often reproduction (while other
individuals are often sterile and don’t reproduce)

Western drywood Honeybee

termite, Incisitermes Apis mellifera
minor queen
Queen, king, soldiers and
workers in termites
 Queens are not genetically
different from workers!
• The question of whether a female individual turns out as a
queen or worker is dependent on the food that she receives as a
larva.
• In honeybees, larvae turn to queens when food nectar contains
more than 35% hexose, and workers when there is less than
10% hexose
• Workers feed “queens to be” with a mandibular gland secretion
called “royal jelly” which ensures that bee queens live for 2-4
years (rather than 7-8 weeks as in workers)
• In addition, the queen is much larger, has bigger ovaries, and
spermatheca
• Can lay 1500 eggs / day!
Characteristics of insect societies
• They live in groups of closely related individuals
• Individuals are usually faithful to their “family”
• Often live in a confined locality (“nest, hive”)
In eusocial insects
• Overlapping generations: parents live with offspring
• Members collaborate in raising the offspring
• Sterility of a large number of individuals of the
colony (e.g. “workers”, “soldiers”): reproductive
division of labour
 Degrees of sociality (after
Michener 1969)
• Solitary (none of the features on previous slide)
• Subsocial: the adults care for their own young for some
period of time (e.g.cockroaches)
• Communal: insects use the same composite nest without
cooperation in brood care (digger bees)
• Quasisocial: use the same nest and also show cooperative
brood care (Euglossine bees)
• Semisocial: in addition to the features in quasisocial, also
has a worker caste (Halictid bees)
• Eusocial: in addition to the features of semisocial, there is
overlap in generations (Honey bees): offspring assist parents
 Size of insect societies
• In honeybees, one queen and up to 40,000 workers
• In bumblebees, one queen and 20-500 workers
• In wasps, 1 to 1000s of queens and 1000s to millions of
workers
• In ants, 1 to over a million queens; up to many million
workers
• One Japanese red wood ant colony contains 307 million
ants, including some 306 million workers and about 1.1
million queens.
• In Hymenoptera, the workers are invariably female; the
males are good only for one thing: sex. Termites also have
male workers
 Success of the eusocial insects
• In the tropics, one third of total biomass
(animal & plant combined!) is made up of
social insects!
• Number of species of social insects
(>11.000) is by far higher than the number
of mammal species (~4500)
Evolution of eusociality in different taxa
Insect order common names frequency of
evolution of
eusociality
Hymenoptera Ants, wasps, bees,
sawflies
11
Isoptera Termites
1
Homoptera Gall-forming
aphids
1
Coleoptera Bark-nesting
weavils
1
Thysanoptera Gall-forming thrips
1
Non-insects Snapping shrimps
and naked mole rats
2
Total 17
A mammal with queen, kings, and a
sterile worker caste
• Naked mole rats
 Reproductive division of labour
• In bees and ants, for example, the queen
typically lays all the eggs
• Sterile individuals (workers, soldiers etc.)
do all the other work, i.e. nest construction,
feeding the young, collecting food, nest
defence etc. – in short, they help the queen
with maximising its fitness, while foregoing
to produce their own offspring.
 Altruistic suicide?
• In many social insects, workers not only forego
reproduction, but also their own life, “for the good
of the colony”
 Altruism and Darwinism?
• The theory of evolution predicts that a trait
(including a behaviour) can only be successful if it
aids its bearer to maximise its fitness (i.e. number
of offspring produced)
• This means that a behaviour should be favoured
by selection if it helps an animal to produce many
copies of the gene that controls the behaviour.
• How can altruism be favoured by natural
selection?
 Why do the workers in social insect
colonies forego reproduction?
• “But with the working ant we have an insect
differing greatly from its parents, yet absolutely
sterile; so that it could never have transmitted
successively acquired modifications of structure or
instinct to its progeny. It may well be asked how is
it possible to reconcile this case with the theory of
natural selection.”
(Charles Darwin)
 William Hamilton’s
concept of inclusive fitness
"I had realized from experience that university people sometimes don't
react well to common sense and in any case most of them listen to it
harder if you first intimidate them with equations.“ Hamilton

• Traditional “direct” fitness is counted as the number of offspring produced.

• Inclusive fitness means that you can still have some fitness if you yourself
produce no offspring – by considering other individuals that share genes with
you.
• For example, you share 50% of genes with your children – hence having two
children will mean, on average, that you have produced one full copy of your
genome. (Hence if you commit altruistic suicide to save more than 2 of your
children, you have won!)
• Any full sibling (brother or sister) shares 50% of your genes, and their children
share 25% of their genes with you.
• Thus, being an uncle or aunt to 4 children is as good as having 2 children of
your own!
• You can increase your own fitness by helping a close relative bring up children!
 Haplodiploid sex determination
in all Hymenoptera

• Haploid male bee copulates with diploid

female -> haploid sperm is stored in
female bee’s spermatheca
• When female “wants” to produce son, she
lays an unfertilised (haploid) egg -> male
offspring
• To produce female offspring, mother
needs to add sperm to egg as it passes
down oviduct
• Only 2 chromosomes shown here
 The queen can decide on the sex of
her offspring!
• In honey bee combs, large cells are for
drones (males) and small ones for workers
(females) – the queen puts only haploid
(unfertilised) eggs into large cells, and only
diploid (fertilised) ones into small cells.
 Fire ant: efficient use of sperm by
the queen
• Like in honeybees, the queen of fire ants
can also actively determine the sex of her
offspring
• On average, only 3 sperm are used to
produce one fertilised egg (female
offspring)
Hymenopteran workers
are more related to their
sisters than to their
(potential) offspring
(Hamilton’s concept of
“supersisters”)
Thus haplo-diploidy implies that
• it might be more beneficial for a worker to help
raise more sisters (workers and new queens) than
to reproduce on her own
• this is because the average relationship between
sisters is higher (0.75) than between workers and
their (possible) offspring (0.5)
• this might explain why the haplodiploid
Hymenoptera have evolved eusocial systems more
than any other insect order
 Coercion – are workers forced
not to reproduce?
• In many species, workers can lay
(unfertilised) eggs, which are haploid and
will turn into males
• Queens “control” the reproduction of their
workers by pheromones, physical
“bullying” and actively eating eggs by
workers
Queen/worker dimorphism allows coercion (Tom Wenseleers)

Bombus terrestris Apis mellifera Nannotrigona melanocera

Vespula vulgaris Atta cephalotes Dorylus wilverthi

 Hamilton’s supersisters idea holds only
if queens are singly mated (i.e. all
workers have only one father)
• but queens of many species (e.g. honeybees,
some ants) mate with multiple partners, and
store all of their sperm
• the result is that there are several “sub-
families” of sisters that are closely related
(i.e. share the same father) in a colony
 Worker policing (F. Ratnieks)
• In cases where queens mate with multiple
fathers, workers often patrol the colony to
ensure that other workers don’t lay eggs
• They eat other worker’s eggs and attack or
immobilise reproductively active females
In two ant species, individuals about to
reproduce are detected by nestmates and
immobilised. In A the queen has
smeared the culprit with a chemical, and
workers physically prevent it from
doing anything at all. In B the “black”
worker hold the “white worker” captive
for days, then turns it over to another
worker for imprisonment
 Altruism in aphids

• Four species of
aphids in which
sterile soldiers
(left) with
enlarged
grasping legs
and short
stabbing beaks
protect their
more delicate
reproducing
colony mates
(right)
 Selfish behaviour in aphid clone
invaders
• Gall-defending aphid
colonies may contain
intruders that have
come from elsewhere.
Although these
outsiders may make up
a substantial proportion
of the group, almost no
newcomers take part in
defence of the gall.
Instead, defenders are
supplied almost
exclusively by the
colony’s native
population (Alcock)
Thrips – a queen (left) and sterile
soldier (right)
 Life cycle of a bumblebee colony
• new queens mate in summer
• all other individuals of colony (old queen, workers, males)
die before autumn
• new queen overwinters and founds new nest in spring
• early in colony development: sterile workers produced
• later in colony development: new queens and males produced
• when colony becomes large and queen old, queen “loses
control”: workers start “rebellion”: attacking queen, and
laying their own eggs
• old queen dies, workers take over reproduction (but produce
only (haploid) males)