 
 
 
 
 
   
between individuals:
 , with
, with 
 
where the number of shared alleles S is sumed over all loci r.
between populations:
 
where the average proportion of shared alleles between and within
populations X and Y (
 ,
, 
 ,
,
 respectively) is computed overall possible combinations of individuals
sampled.
respectively) is computed overall possible combinations of individuals
sampled.