2
Null model analysis has been a popular tool for detecting pattern in binary presence–absence matrices, and previous tests
have identiied algorithms and metrics that have good statistical properties. However, the behavior of diferent metrics
is often correlated, making it diicult to distinguish diferent patterns. We compared the performance of a suite of null
models and metrics that have been proposed to measure patterns of segregation, aggregation, nestedness, coherence, and
species turnover. We found that any matrix with segregated species pairs can be re-ordered to highlight aggregated pairs. As
a consequence, the same null model can identify a single matrix as being simultaneously aggregated, segregated or nested.
hese results cast doubt on previous conclusions of matrix-wide species segregation based on the C-score and the ixed-ixed
algorithm. Similarly, we found that recently proposed classiication schemes based on patterns of coherence, nestedness,
and segregation and aggregation cannot be uniquely distinguished using proposed metrics and null model algorithms. It
may be necessary to use a combination of diferent metrics and to decompose matrix-wide patterns into those of individual
pairs of species or pairs of sites to pinpoint the sources of non-randomness.
Pattern detection in null model analysis
Werner Ulrich and Nicholas J. Gotelli
W. Ulrich (ulrichw@umk.pl), Nicolaus Copernicus Univ. in Toruń, Chair of Ecology and Biogeography, Gagarina 9, PL-87-100 Toruń,
Poland. – N. J. Gotelli, Dept of Biology, Univ. of Vermont, Burlington, VT 05405, USA.
Oikos 122: 2–18, 2013
doi: 10.1111/j.1600-0706.2012.20325.x
© 2012 he Authors. Oikos © 2012 Nordic Society Oikos
Subject Editor: Paulo Guimaraes Jr. Accepted 16 March 2012
C
h
o
i
c
e
E
d
i
t
o
r
’
s
OIKOS
A major research efort in community ecology has been
to infer mechanisms of community organization from
patterns in a binary presence–absence matrix. In such a
matrix, species are represented by rows, sites are represented
by columns, and the entries are the presence or absence of
a species in a particular site (McCoy and Heck 1987). Early
research (and controversy) focused on the role of inter-
speciic competition in producing checkerboard patterns
(Diamond 1975), species segregation (Stone and Roberts
1990), and missing species combinations (Pielou and Pielou
1968). A largely independent focus on species nestedness
(Darlington 1957, Patterson and Atmar 1986) emphasized
the role of selective, orderly extinction (and immigration)
in producing species-poor assemblages whose composition
is a perfectly nested subset of more species-rich assemblages.
Leibold and Mikkelson (2002) signiicantly expanded
this framework by proposing tests for additional patterns
of Clementsian, Gleasonian, and evenly-spaced gradients.
heir method involves re-ordering the rows and columns
of a presence–absence matrix by reciprocal averaging to
maximize the coherence of species range sizes. A maxi-
mally coherent matrix state is one that contains the greatest
number of uninterrupted sequences of species occurrences
within rows of the re-ordered matrix. In such a re-ordered
matrix, ‘species turnover’ is the replacement of one species
by another, and ‘boundary clumping’ is the extent to which
species boundaries are clumped or over-dispersed. Checker-
boards, nestedness, Clementsian gradients, Gleasonian
gradients, and evenly-spaced gradients are recognized by a
combination of patterns of coherence, species turnover, and
boundary clumping.
Presley et al. (2010) further expanded Leibold and
Mikkelson’s (2002) framework, distinguishing among
12 diferent models based on the combination of bound-
ary clumping and turnover patterns for matrices in which
there is positive coherence. hey distinguished three types
of nested patterns, depending on whether boundary clump-
ing was positive, random, or negative, and they recognized
six additional ‘quasi-structures’ based on weakly positive or
negative patterns of species turnover.
he review and decision to publish this paper has been taken by the above noted SE.
he decision by the handling SE is shared by a second SE and the EiC.
Synthesis
The identification of distinctive patterns in species x site presence-absence matrices is important for under-
standing meta-community organisation. We compared the performance of a suite of null models and metrics
that have been proposed to measure patterns of segregation, aggregation, nestedness, coherence, and species
turnover. We found that any matrix with segregated species pairs can be re-ordered to highlight aggregated
pairs, indicating that these seemingly opposite patterns are closely related. Recently proposed classification
schemes failed to correctly classify realistic matrices that included multiple co-occurrence structures. We
propose using a combination of metrics and decomposing matrix-wide patterns into those of individual pairs
of species and sites to pinpoint sources of non-randomness.