Conclusion - An Introduction to Behavioural Ecology - page 431

Wynne-Edwards, would be
when this also coincides
with inclusive fitness. This
requires extremely restrictive
conditions, with extremely
high relatedness within groups
(e.g. clonality, r = 1) or some
mechanism such as policing
that completely aligns the
interest of the individual and
the group (Gardner & Grafen,
2009).
Although this alignment of
inclusive and group fitness
should happen extremely
rarely, when it does it can have
huge consequences because it
can lead to a major
evolutionary transition. A
major evolutionary transition
is when groups of individuals
that were previously capable of
independent reproduction
before the transition can only
replicate as a larger unit after it
(Maynard Smith & Szathmary,
1995; Fig. 15.2). In Chapter
13 we discussed the transition
to eusociality, where colonies
of social insects have become
'superorganisms' - this
represents one of only
approximately eight major
evolutionary transitions which
have occurred (Table 15.2).
The major transitions
approach makes it clear that
the behavioural ecology
problem of cooperation has
played a central role in
evolutionary progress, because the problem of cooperation must be solved if a group
of individuals is to come together to form a new, more complex, organism.
Furthermore, similar to the examples discussed in Chapter 12, the major evolutionary
transitions can be divided between those which have involved either relatives (e.g. the
evolution of multicellularity and eusociality) or non-relatives (e.g. the evolution of
chromosomes and the eukaryotic cell; Queller, 2000), and hence either indirect or
direct benefits to cooperation.

The alignment of
inclusive and
group fitness can
lead to a major
evolutionary
transition

Fig. 15.2 Thomas Hobbes' Leviathan shows a
giant crowned figure, which is composed from
over 300 humans. The quote above is from the
Book of Job and translates as 'There is no power
on earth to be compared with him'. However, the
major transition approach emphasizes that this is
exactly what happens on earth, with groups of
individuals coming together to form higher level
individuals.