Cooperative Breeding

Many animals provide care for young that are not their own direct
offspring. This is often accompanied by a delay in or surrender of
their own reproduction.
This type of behaviour often has been called altruism because there is an
apparent cost to the giver and an apparent benefit to the receiver. How can
this work in the context of natural selection?

Florida Scrub Jays helping at the nest

Heritable traits, natural selection and
evolution
A fundamental question; Is the giver of the behaviour “willing”
or constrained in some way? Constraints may be;
•Ecological (lack of other options), or
•Behavioural (coercion by dominants)
A closer look at these situations reveals that in nearly all
examples in which cooperative breeding exists, there is also an
extended family structure or kin group.

Remember
def’n of
parental
care!

N3 is number of EXTRA young raised by parents as a result of help from non-parents

r = chance that an allele in one individual is present in another because of
inheritance from a common, recent ancestor – what we call common descent

(of parent’s genes)

(of helper’s genes)

Inclusive
Fitness is the
average sum
total of alleles
Coefficients of relatedness (r) between relatives (normally)
RELATIONSHIP r
Full siblings 0.5 These are averages!
Parent-offspring 0.5
Meerkats are cute and social,
Half-siblings 0.25 but somewhat selfish!
Uncle/aunt-niece/nephew 0.25
Cousin-cousin 0.125
Grandparent-grandchild 0.25
The strength of selection to help
kin is expected to be, on
average, proportional to “r”,
thus one expects to see closest
kin helping each other most,
although this is enormously
variable
 If an animal increases the # of non-descendant kin surviving
by helping in some way, and, if this gain in indirect fitness is >
than the loss in direct fitness by not breeding, then genes for
helping behaviour can spread through a population. This is
Hamilton's Rule. r b B > rcC

Indirect fitness
also may be
increased by Exp = helpers removed
increasing the
probability of
survival of the
parents if helping
does this, because
they then
produce more
kin.
Benefits of helping
in the Florida
Scrub Jay
Helpers often also gain by an increased chance of becoming
breeders themselves at a later date--either by inheriting a
territory or by helping to enlarge it to the point where extra
breeding is possible. Where going it alone is not possible because
of distribution and number of suitable breeding territories,
delaying and cooperation are likely to evolve.
Sociality comes in different forms
The issue of living together for cooperative breeding is one
largely of the advantages of communal living versus the costs
of going it alone; as well as benefits of kin selection working
together to maintain social groups—especially family groups.

“Related” helper Pied

Kingfishers deliver more
food to nest than
“unrelated” helpers.
Delayed breeding, often in cooperative family units is expected
to occur when;
•The production of mature offspring exceeds the availability of
acceptable opportunities for independent reproduction

Hamilton’s Rule; rbB>rcC

Under these circumstances, offspring must postpone breeding until an

opportunity presents itself and they are capable of successfully competing
And, in these circumstances, one might as well wait at home where helping
provides indirect fitness at least
Advantages of sociality:
•Many eyes to spot predators
•Mobbing/cooperation in response to predators
•Greater ability to hold territories
Disadvantages; for sociality
•Greater competition for food, nesting sites etc-conspecifics are
expected to compete most intensely
•Greater transmission of parasites from animal to animal and
nest to nest
•More difficult to be cryptic re predators
Sociality exists in a number of grades;
Males and females that are normally solitary aggregate for
mating
Bachelor herds of male elephants have loose associations

Females may also

associate--often
mother/daughter
relationships in extended
families
Eusociality
Eusociality (bees, wasps, ants, termites, aphids, naked mole
rats) is the most extreme form of social living where some
members of colony do not produce their own offspring (but
help rear lots of siblings).
Characterized by:
Long lived colony with 2 or more overlapping generations
Cooperation in maintaining colony and raising young of
usually a single primary reproductive

Obligatory roles (behaviours)

and often correspondingly
polymorphic in appearance
(castes)
Haplodiploid Genetics
Hamilton’s Rule;
rbB>rcC
A closer look at these situations reveals that in nearly all
examples in which cooperative breeding exists, there is also an
extended family structure or kin group.

Remember
def’n of
parental
care!

N3 is number of EXTRA young raised by parents as a result of help from non-parents

r = chance that an allele in one individual is present in another because of
inheritance from a common, recent ancestor – what we call common descent

(of parent’s genes)

(of helper’s genes)

Inclusive
Fitness is the
average sum
total of alleles
Inbreeding
Where inbreeding has occurred for a long time so that
deleterious homozygotes have largely been selected out, the
result may be a very homogeneous genome
When inbred males mate inbred females, offspring share
essentially 100% of genome-->> 50% for average parents and
offspring

Heterocephalus glaber