158
1
1
DT HHT
z
z
z
PP k z
N
kk
bg()=-expl(),,a + og lo
æ
è
ç
ö
ø
÷
æ
è
ç
ö
ø
÷+
=
+
++
+
++
å gg,N
é
ë
ê
ù
û
ú
(10a)
where HP,γ and HP,α denote respectively the gamma and alpha phylogenetic entropy.
When the species importance measure zik represents the ith species relative abun-
dance in the kth current-time assemblage, then zz++kk== 11 ,,++Nz//zN++=. In
this special case, we have^1 DTbg()=-exp/ëé()HHPP,,a Tùû. Thus an additive
decomposition for phylogenetic entropy HP holds (Pavoine et al. 2009 ; Mouchet
and Mouillot 2011 ), as for ordinary Shannon entropy (Jost 2007 ).
(c) When q = 2, the phylogenetic beta diversity can be expressed as
2 1
2
2
DT
Lz
Lz
T
T
N
i
k
N
ik
ii
i
b()= N.
Î=
+
Î
åå
å
iB
B
In the special case of zz++k==1, + N, this phylogenetic beta diversity of order 2
can be linked to quadratic entropy as
(^2) DT 11 QT^1 QT^1
bg()=-()//()- a/,
- (10b)
where Qγ and Qα denote respectively the gamma and alpha quadratic entropy. The
above formula is also applicable to non-ultrametric trees by replacing all T with T,
the mean branch length in the pooled assemblage; see Chiu et al. ( 2014 , Appendix
C) for a proof.
Normalized Phylogenetic Similarity Measures
For traditional abundance-based diversity, the most commonly used similarity mea-
sures include N-assemblage generalizations of the Jaccard et al. ( 1966 ) and Morisita-
Horn (Morisita 1959 ) measures. The latter three measures were integrated into a
class of CqN measures by Chao et al. ( 2008 ). Jost ( 2006 , 2007 ), Chao et al. ( 2008 ,
2012 ), and Chiu et al. ( 2014 ) have demonstrated that all the above measures are
monotonic transformations of beta diversity based on the ordinary Hill numbers.
This is an advantage of using the framework of Hill numbers: a direct link exists
between diversity and similarity (or differentiation) among assemblages.
Chiu et al. ( 2014 ) extended this framework by proposing four classes of similar-
ity (or differentiation) measures that are monotonic functions of phylogenetic beta
diversity. The basic idea is that the phylogenetic beta diversity, a ratio of gamma and
alpha phylogenetic Hill numbers, is independent of alpha and measures the pure
differentiation among assemblages. The phylogenetic beta component always lies
A. Chao et al.