Social and Eusocial Behavior

Why do animals help others at the

potential cost of their own survival
and reproduction?
Alka Nokhwal
Ph.D., M.Sc. (Zoology)
Senior Research Fellow
NCVTC, ICAR-National Research Centre on Equines,
Hisar, Haryana
Social behavior consists of a set of interactions among
individuals of the same species.

Highly social
organisms live
together in large
groups, and often
cooperate to conduct
many tasks (Ex. packs
of wolves and schools The most highly
Some animals rarely if
of fish). social animals form
ever interact with one
another, even when it tightly knit
comes to issues of colonies(Ex. all ants
parental care (Ex. and termites, some
mosquitoes and polar bees and wasps
bears).
etc.).

Wide
range of
sociality
 Social Behavior is Adaptive

Many social behaviors of animals are adaptive,

meaning that being social ultimately increases an
animal’s fitness — its lifetime reproductive success.
 One example of how social behavior is adaptive is
aggregation against predators. This concept applies to
caterpillars feeding together on a leaf, a herd of
wildebeest, schools of fish, and flocks of birds.
 Wildebeests gathered into groups are more protected from predators than any
solitary wildebeest.
 Costs and Benefits

• Living in groups • Benefits exceed

the costs and
risks of social
life
Social cooperation
Balance of conflict favored
and cooperation
 Altruism

An altruistic act is one that increases the welfare of another individual

at an actual or potential cost of the individual who performs the act.

The benefit of an altruistic behavior is ultimately measured in its

effect on an animal’s lifetime reproductive success. Evolutionary
biologists and animal behaviorists have sought to identify the
mechanisms that can explain what some have called the “problem of
altruism.” Natural selection operates against individuals who reduce
their own fitness. Altruism by definition decreases the fitness of the
individual, so how can this behavior persist?
 benefits of
altruism can
existence of
Reciprocity outweigh the
altruism
costs of altruism
in long term
• Eusociality
The evolution of social behavior at its most intimate and complex
degree is found in eusocial animals. Eusocial species live in
colonies. Only a relatively small fraction of the animals in the
colony reproduce; the non-reproductive colony members provide
resources, defense, and collective care of the young.
 The list of known eusocial animals includes ants, termites, some
wasps, some bees, a small number of aphid and thrip species, two
species of mammal (the naked mole rat and the Damaraland mole
rat), and multiple species of reef-dwelling shrimp.
Eusocial
behaviour
Eusocial animals share the following four
characteristics:

Adults live in groups,

Cooperative care of juveniles

(individuals care for brood that is
not their own),

Reproductive division of labor (not

all individuals get to reproduce)

Overlap of generations
Other types of social interactions

• Social behavior between

Subsociality parents and offspring (e.g.,
birds, Halictine bees

Parasocialit • Social behavior among

y members of the same
generation (e.g., most bees)
Which Animals are Eusocial?
• Termites are thought to be highly developed social insects that live
inside the food they consume (rotting wood; Thorne 1997). Termites
are diploid insects that engage in intricate social behaviors like nest
building and territorial defense.
• most recent eusocial organisms to be found- shrimp, aphids, and
thrips.
• The Synalpheus shrimps' eusociality has at least two distinct origins.
As internal parasites on tropical sponges, these marine shrimp thrive
in colonies of several hundred closely related diploid individuals.
Given that dispersing to found new colonies is riskier than remaining
in the natal nest, the variable distribution of the host sponges may
have aided in the evolution of eusociality within this group.
• Thrips are small haplodiploid insects in the order
Thysanoptera. 300 of the approximately 5,000 species
build nests in plants called galls where they feed on the
plant tissue. Of these, six species can be categorized as
being eusocial since they have militaries with varying
morphologies that protect the galls from kleptoparasites.
• Like thrips, social aphids live in plant galls or hollow
stems and feed on plant tissue. These tiny hemipterans can
breed parthenogenetically while having diploid life cycles,
and several species have been described with strong soldier
morphs.
 There are at least two species of vertebrates that
could be considered eusocial, the naked mole
rat and the Damaraland mole rat.
• Both species are diploid, highly inbred and live
in harsh deserts with patchy food resources.
• Most individuals help to raise siblings or close
relatives that are born to a single reproductive
female (the queen).
 Advantages/Disadvantages to Living in
Groups

Groups may form as Acquiring food in

Advantages against
defense against
competitors, e.g., ant groups (e.g.,
predation, forming a
"selfish herd"
Azteca trigona raiding army ants;
(Adams 1994). Solé et al. 2000).
(Hamilton 1971).

Increased Easy detection of

Increased transmittance of the group by
competition parasites and predators and
diseases parasites.
 How did Eusociality Evolve?
• Natural selection's core element is in contrast with giving up
one's capacity for reproduction (to survive and reproduce).
• According to evolutionary biologists, the development of
eusociality followed a path that began with solitary organisms
learning the advantages of group behaviour and eventually
reached a "point of no return" (Wilson & Hölldobler 2005)
where some individuals lost their ability to physically reproduce
and could only benefit indirectly from evolution.
• It's also important to keep in mind that the selecting pressures at
play at the time eusocial behaviour first emerges may not be the
same as those sustaining advanced eusocial colonies (Hölldobler
& Wilson 2009).
 Ecological
contributions

Kin selection
Primitive Point of Advanced
Eusociality “No return” eusociality
Delayed
benefits

Multi-level
selection

It is important to note that they are not mutually exclusive - each

may play a different role in the evolution of eusociality in
different groups.
 Kin Selection
• A gene can spread copies of itself in subsequent
generations either directly by producing
offspring or indirectly by promoting the
reproduction of close relatives.
• Inclusive fitness is the total of all reproductive
benefits, both direct and indirect. As a result, if
indirect fitness levels are higher than direct
fitness, eusociality may be chosen over solitary
behaviour.
• An altruistic act is one that helps the recipient at the expense of the performer.
• According to Hamilton's rule (Hamilton 1964), altruism is preferred if
r > C/B,
• where C is the cost and B is the gain to the recipient of the altruistic act in terms of
lifetime reproductive success (decrease in lifetime reproductive success). The percentage
of alleles shared by two people who are related by ancestry is known as the coefficient of
relatedness, or r, and it extends from 0 to 1. High degrees of altruism within groups are
necessary for eusociality.
• Inbreeding and haplodiploid sex determination are two well-known ways that might
increase ‘r’.
• Hamilton's rule is weighted in favour of raising sisters rather than kids in haplodiploid
organisms because the relatedness between full sibling sisters (r = 0.75) is higher than the
relatedness between a mother and her offspring (r = 0.5).Inbreeding produces offspring
that share a greater percentage of alleles, raising r. This is typical of species that don't
travel very far from their natal nest or are more likely to mate with their siblings (e.g.,
termites and wild naked mole rats).
 Delayed Benefits
• "Hopeful reproductives" — workers with the option to
stay and assist or go away and start their own nest — are a
possible intermediate step toward eusociality.
• The choice may be influenced by factors like territory, food
availability, environmental factors, and group hierarchy.
Until there is a chance to take over as parents, Florida scrub
jay young are known to remain at the natal nest, raising
siblings and improving their inclusive fitness.
• Primitive eusocial wasp colonies, like Polistes, are
frequently passed over to dominant workers after a queen
dies.
 Multi-level Selection
• Natural selection can happen on an individual, family (a group of
related people known as "kin"), or group level (non-related
individuals).
• Models of multi-level or trait-group selection can be used to
illustrate how the traits (phenotype) of the colony interact with
the environment to determine colony-level fitness in eusocial
organisms.
• It is still up for debate whether models of multi-level selection or
inclusive fitness models are the best way to investigate the
development and maintenance of eusociality, particularly given
the paucity of empirical data supporting inclusive fitness in
groups (Seeley 1997).
Ecological and Life History Contributions

• Nesting behaviour has been suggested as a potential precondition for

the growth of eusociality, in part because it fosters circumstances that
encourage cooperative brood care (Anderson 1984). "Fortress
defenders" can work together to protect this vital resource in areas
where nest founding is hazardous or there are few territories or spaces.
• Parental care can also play a significant role in a person's life story.
One route to eusociality in Hymenoptera is believed to begin with
solitary females participating in concurrent progressive provisioning,
which involves raising several larvae of various ages at once.
• The next step in the transition to eusocial behaviour would be for the
surviving offspring and provisioning siblings, then for the offspring to
refrain from having children of their own.
Thank you