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Austral Ecology (2012) 37, 903914

Assembly rules operate only in equilibrium communities:

Is it true? aec_2349 903..914

FLORENCE B. J. CASH,1 ARRON CONN,1 SEAN COUTTS,1 MELANIE STEPHEN,1

NORMAN W. H. MASON,2 BARBARA J. ANDERSON3 AND J. BASTOW WILSON1*
1
Botany Department, University of Otago, PO Box 56, Dunedin 9054, New Zealand (Email:
bastow@bastow.ac.nz), 2Landcare Research, Hamilton, New Zealand; and 3Biology Department,
University of York,York, UK

Abstract Very little is known of how disturbance affects community assembly rules. We examine this in three
disturbance states in each of two ski areas on southern New Zealand mountains.Theory suggests that a community
will become progressively more spatially organized during recovery from disturbance. Firstly, different patches of
the community should become more similar through time, but this was seen in only one of the two areas and even
then only examining species presence/absence. Secondly, it has been suggested that spatial autocorrelation will be
stronger in less-disturbed conditions, that is, there will be a stronger pattern of more distant patches being more
dissimilar in species composition. This was generally borne out. However, the method indicated more point
randomness in less-disturbed sites. Assembly rules might be seen in species abundances. Previous work has found
maximum evenness of abundances in later successional communities, but the pattern here was the opposite: high
evenness in the most disturbed communities. The literature suggests that in undisturbed communities the distri-
bution of species abundances (relative abundance distribution) will be general lognormal, and we further argue that
the identity of the species across occupying rank positions in that distribution should be more consistent (rank
consistency). Both predictions were borne out in one area, but neither in the other. Many workers suggest that
niche-based assembly rules will be stronger in undisturbed communities. However, there was only weak evidence
of constancy in species richness. Local species assemblages tended to contain a relatively constant representation
from different morphological/taxonomic guilds (guild proportionality) and this was significant in some tests, but
contrary to theory this effect occurred mainly in the most disturbed sites. It is concluded that there is only limited
truth in the frequent assumption that community structure is stronger in undisturbed, equilibrium communities.

Key words: community structure, distance decay, disturbance, heterogeneity, species abundance distribution.

INTRODUCTION one or other species or groups of species (not simply

An understanding of the structure of ecological com-
munities of spatial heterogeneity and of species
assembly rules is necessary background for work in
the conservation of biodiversity, in prediction of com-
munity resilience and in setting the targets for ecologi-
cal restoration. However, communities are not static;
disturbance is endemic to all of them, and we need to
know how heterogeneity and assembly respond to
disturbance. These are questions of both theoretical
and practical importance.
Initial random colonization might lead to spatial
disorder just after a disturbance, but after an interme-
diate stage competitive sorting could lead to spatial
segregation later (Gitay & Wilson 1995). Disturbance
may also affect assembly rules, defined as restrictions
on the observed patterns of species presence or abun-
dance that are based on the presence or abundance of
the response of individual species to the environment)
(Wilson 1999, cf. 2007). Soon after a disturbance,
density dependence might not yet have been restored,
inferior competitors might not have been competi-
tively excluded, and the relative abundances of species
might still be adjusting to dispersal, competition, etc.
Many authors have argued that since assembly rules
are the result of these processes they might be absent
then, that is, the assembly rules will not be seen until
later in succession as communities approach equilib-
rium (e.g. Belyea & Lancaster 1999; Drake et al. 1999;
Stokes & Archer 2010). We can interpret this in terms
of spatial community assembly, abundance-pattern
assembly rules and niche-based assembly rules.
Spatial community assembly

*Corresponding author. We first examine spatial heterogeneity, that is, the

Accepted for publication December 2011. mean point-to-point variation in species composition
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904 F. B. J. C A S H ET AL.
within the community. Prediction from theory here is
complex (Gitay & Wilson 1995). Early in succession,
heterogeneity might be high due to chance
colonization. This effect would decrease through time,
but heterogeneity might rise eventually due to sorting
of the species into communities (Childress et al.
1998). If there is a positive-feedback switch, in which
the species in each patch modify their environment to
their own advantage, giving alternative stable states,
the later increase in heterogeneity in species composi-
tion and micro-environment would be further rein-
forced (Wilson & Agnew 1992; Mason et al. 2007).
Chaos theory predicts a similar result, suggesting that
initial heterogeneity will sometimes be magnified with
time (Stone & Ezrati 1996). Thus, theory indicates
higher heterogeneity just after severe disturbance, and
then again in undisturbed communities.
An additional aspect of heterogeneity is the spatial
trend, that is, spatial autocorrelation. At the commu-
nity level this is the ability to predict the dissimilarity
of two patches of vegetation from their distance apart
(Mistral et al. 2000). The strength of this relation is a
measure of the communitys spatial structure, and
logic suggests that so long as the focus (e.g. quadrat
size) is smaller than the grain of micro-environmental
variation, predictability should increase with time from
disturbance, as species composition responds to varia-
tion in micro-environmental conditions. However,
there is only limited and contradictory evidence on
this (e.g. Seabloom et al. 2005; Erfanzadeh et al.
2010).
A spatial autocorrelation fit can also be extrapolated
backwards to zero distance (the nugget), at which all
environmental dissimilarity and dispersal limitation
have been removed, giving an estimate of the point
randomness of the community (Fortin & Dale 2005;
Lawrence Lodge et al. 2007). We argued above that
this should be higher in more disturbed communities.
Thus, the spatial community assembly hypotheses
are that:
1. Spatial heterogeneity: will be high in the most
disturbed community and in undisturbed commu-
nities, but lower in those with intermediate
disturbance.
2. Spatial autocorrelation strength: will be greater in
the least disturbed communities.
3. Spatial autocorrelation nugget: will be smaller in
the least disturbed communities.
Abundance-pattern assembly rules
Community re-assembly after a disturbance involves
adjustment of the abundances of the species, and the
most basic aspect of this is evenness (Bartha et al.
2001).There is little theory to guide us on the response
here, but the most rigorous tests to date have shown
doi:10.1111/j.1442-9993.2011.02349.x
evenness to increase through succession, at least in
herbaceous/shrubby communities (Wilson et al. 1996).
To analyse abundances more closely, we examine
relative abundance distributions (RAD), the
dominance/diversity relations of early workers. These
have also been related to succession, with the sugges-
tion from mathematical modelling that a General Log-
normal distribution will be seen in equilibrium, late-
successional communities but not in disturbed ones
(Stenseth 1979; Ugland & Gray 1982). However, this
remains largely unexplored, especially for plant
communities.
Ecologists have distinguished between ecological/
founder control, whereby composition depends on the
chance order of species arrival, versus evolutionary/
competition control, whereby it depends on the
(evolved) competitive abilities of the species. This will
be reflected in whether the ranks of an RAD are occu-
pied by the same species in different patches/quadrats,
for example, whether the most abundant species is
always has the same identity (Watkins & Wilson 1994).
Just after a disturbance, the ranks should depend
largely on chance dispersal (i.e. ecological/founder
control), giving low consistency between patches/
quadrats. As the community assembles, rank consis-
tency should increase as the relative competitive
abilities of species determine their presence and abun-
dance (evolutionary/competition control).
Thus, the hypotheses for abundance-pattern assem-
bly rules are that:
4. Evenness: will be highest in the undisturbed
community.
5. Relative abundance distribution: the undisturbed
communities will show a lognormal RAD; com-
munities with less disturbance and especially the
most disturbed will show geometric, Zipf-
Mandelbrot or possibly broken stick RADs.
6. Rank consistency: will be higher the less disturbed
the community, that is, there should be more con-
sistency in the identity of species in the ranks of
the RAD.
Niche-based assembly rules
The concept of the niche (Hutchinson 1957) implies
that, in the longer term, limiting similarity (MacArthur
& Levins 1967) will restrict the number of species to
the number of distinct niches: niche limitation. The
identity of the species may be different in different
patches of a community because of chance coloniza-
tion and subtle micro-environmental variation, but
there should be a trend towards less variation in
species richness, that is, greater constancy in species
richness, than if the species occurred independently of
each other (Wilson et al. 1987). The appropriate null
model for such independent occurrence of species
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fixes the number of occurrences of each species at that
observed in the real data, but assigns those occur-
rences at random to sub-quadrats, independent of the
assignment of other species (Wilson et al. 1987).
Likewise, it should be easier for a species to invade
or to avoid exclusion if it is in a different guild from the
majority of the species already present so long as the
guilds represent the limitations to co-existence. This
would give a more constant representation of different
guilds than under a null model which randomizes
occurrences while retaining species frequencies and
quadrat richness: the assembly rule of guild propor-
tionality (Wilson 1989). We here use a priori guilds,
based on morphology/taxonomy.
However, after disturbance there might be dispersal
limitation, empty niches (Hutchinson 1959) and
incomplete competitive exclusion, so these rules would
not apply. Thus, the niche-based assembly rule
hypotheses are that:
7. Constancy in species richness: in less-disturbed
communities, species richness will show lower
variance between sub-quadrats than expected at
random.
8. Guild proportionality: in less-disturbed communi-
ties, the proportions of species from each guild
will show lower variance between sub-quadrats
than expected at random.
study of human impacts. To summarize, we examined
the eight hypotheses listed above, under the following
categories:
Spatial community assembly:
1. spatial heterogeneity,
2. spatial autocorrelation strength,
3. spatial autocorrelation nugget,
Abundance-pattern assembly rules:
4. evenness,
5. relative abundance distribution,
6. rank consistency,
Niche-based assembly rules:
7. constancy in species richness, and
8. guild proportionality.
METHODS
Sites
Two areas with extensive ski activity were chosen on nearby
mountain ranges in southern New Zealand (Fig. 1). The
Cardrona area (4552S, 16856E) was at 15601640 m
a.s.l., with 2.6 m per year of snowfall, and Coronet Peak

Purpose

In spite of all this theory, there is actually little evi-

dence that there is more spatial disorder or weaker
assembly rules after disturbance. Wilson et al. (2000)
did find that in two semi-arid grassland sites, the one
which had been more disturbed gave no evidence that
species constrained each others biomass, whereas the
less-disturbed site did. Sanders et al. (2003) found that
disturbance of an ant community in the form of inva-
sion by an exotic species reduced the chequerboarding
pattern, which they saw as breakdown in assembly
rules. However, there seem to have been no direct
comparisons of assembly rules under various degrees
of disturbance, or of the differences in spatial organi-
zation that would be expected to follow.
We examined the relation between disturbance,
spatial community assembly and assembly rules using
ski-related disturbance on two southern New Zealand
mountains (a minimal form of replication). Distur-
bance of ecological communities generally comprises
either repeated removal of vegetation (usually mowing
or grazing) or mechanical disturbance exposing bare
soil (e.g. by burrowing animals). Ski-slope making
combines the two. Mechanical soil disturbance is one
of the major factors involved in human perturbation of
plant communities, so determining its effect on assem- Fig. 1. South Island, New Zealand. Location of the two
bly rules during ski-run formation is a significant case areas sampled.

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906 F. B. J. C A S H ET AL.
(4556S 16843E, referred to as Coronet below) at 1180 was recorded. In the first row of 10 sub-quadrats in each
1200 m a.s.l., with extensive snowmaking. Within each area,
three sites were chosen:
Ski-run
These runs were created at Cardrona in 19781979 and at
Coronet in 1947. At intervals of about 5 years, rocks are
removed, on some occasions the vegetation is removed and
then replaced but on other occasions a mixture of alien
species is sown. Disturbance is caused by the machinery
brought in for this. At Cardrona the sown mix comprised
Agrostis capillaris, Festuca rubra and Trifolium repens, at
Coronet only A. capillaris. The work is carried out every few
years about April (autumn), with sowing timed for germina-
tion the following spring. Sampling was conducted in April,
but the sites sampled had not been disturbed for at least
3 years. A few native and alien forbs had colonized. All these
species clearly arrived by seed. Native Poa colensoi was
present on the ski-run at Cardrona, but it too commonly
invades by seed in alpine areas in the region (Roxburgh et al.
1988).
Adjacent
These sites were near the ski-runs, somewhat disturbed in the
making and maintenance of the ski-run. The vegetation was
mainly native, with species such as Chionochloa rigida (snow
tussock) and subshrubs Gaultheria depressa and Dracophyllum
pronum. These are very likely to be remnants of the pre-
disturbance vegetation. Poa colensoi could have invaded since
disturbance, but since it is no more frequent than in the
Undisturbed sites it has probably remained. Small amounts
of alien grasses were present: Agrostis capillaris at both sites,
with Anthoxanthum odoratum at Cardrona and F. rubra at
Coronet.
Undisturbed
These comprised tussock grassland undisturbed by ski
activity. Although labelled Undisturbed for brevity, the
original shrub cover had probably been reduced by pastoral
burning before 1970, but there were no current signs of
disturbance. Species composition was similar to the respec-
tive Adjacent sites.
In terms of our hypotheses, the degree of disturbance is
not precisely defined, but the two extremes above seem to
match clearly highly/recently disturbed and undisturbed
respectively.
Sampling
At each of the three sites in each area, six 1 1 m quadrats
were placed by restricted randomization. Each quadrat was
divided into a grid of 100 sub-quadrats, each 10 10 cm, in
which the shoot presence/absence of all vascular plant species
doi:10.1111/j.1442-9993.2011.02349.x
quadrat, all above-ground plant material in the volume
defined by each sub-quadrat was collected, sorted by species,
and the biomass determined.
Analyses: spatial community assembly
To measure spatial heterogeneity in the community at a site,
the mean dissimilarity was calculated between all possible
pairs of 10 10 cm sub-quadrats within each quadrat.
Within each of the six quadrats, 4950 pairs were available for
presence/absence, 45 for biomass. The dissimilarity measure
for the grid of presence/absence records was Jaccard (1.0
Jaccard similarity: Jongman et al. 1987), and for the line of
biomass records proportional dissimilarity (PD: Jongman
et al. 1987). Jaccard and PD both give 0.0 for identical
species composition and 1.0 for maximum dissimilarity,
and neither is affected by species absent from both samples
being compared. These values were used in a two-way
disturbance area analysis of variance, with the six quadrats
within each site used as replicates.
For spatial autocorrelation within a quadrat, the same
dissimilarities were compared with the distance apart of the
two sub-quadrats by fitting the asymptotic relation (cf.
Lawrence Lodge et al. 2007):
Distance
Dissimilarity = nugget + b
c + Distance
The dissimilarities and distances within each of the six
quadrats were accumulated for each site in each area. The
goodness of fit (% of the variance dissimilarity explained by
distance using this formula) for each quadrat was arcsine
transformed and used in analysis of variance (an alternative
logit transformation gave the same pattern of significances).
Transformation removed minor heteroscedasticity, but the
means and analysis results were very similar to those on
untransformed data and untransformed means are
presented. The intercept is termed the nugget in spatial
autocorrelation, has been seen as an estimate of community
randomness (Fortin & Dale 2005). The significance of the
relation was examined by a randomization test in which the
species composition of a sub-quadrat was kept as a unit, but
randomized 2000 times with respect to the position of the
sub-quadrat within its quadrat (Lawrence Lodge et al. 2007).
Analyses: abundance-pattern assembly rules
Evenness in species biomass was measured in each of the 10
biomass sub-quadrats of a quadrat using the Camargo index
E, selected for its ability to meet the 15 requirements and
features evaluated by Smith and Wilson (1996). The means
over the sub-quadrats were used in analysis of variance.
To characterize the RAD, four models of RAD (Broken
Stick, Geometric, General Lognormal and Zipf-Mandelbrot)
were fitted to each of the 10 10 cm biomass sub-quadrats
by the method of Wilson (1991). In order to allow for any
tendency for all models to fit some observed distributions
poorly, that is, to compare models on a standard basis, the
sum of squares of deviations from the best-fitting model was
subtracted from all four models, separately for each sub-
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quadrat (Wilson et al. 1996), and quadrat means calculated
over these sub-quadrat values.
Rank consistency across the ten biomass sub-quadrats of a
quadrat was determined with index Cr (Watkins & Wilson
1994), which ranges from +1 when the ranks of the species (1
if the species is the most abundant, 2 if it is the second-most
abundant, etc.) are identical between replicate sub-quadrats
and -1 when they are as different as they could be. The null
model used randomly permutes the non-zero species abun-
dances within a sub-quadrat. The observed value of Cr was
compared with random expectation using an IV index:
Observed
IV = 200 100
Observed + Expected
where Observed and Expected are, respectively, the values of
the test statistic (here, Cr) observed and that expected under
the null model (i.e. the mean of randomizations). Such an
index takes a positive value (>0, to a maximum of +100)
when the observed value is greater than expected under the
appropriate null model (see below), and a negative value (<0,
to a minimum of -100) when the observed value is less than
expected under the null model (Watkins & Wilson 1994).The
expected values were the means over 5000 randomizations.
These IV values, one per quadrat, were used in analysis of
variance.
Analyses: niche-based assembly rules
For these analyses, constancy in species richness and guild
proportionality, the test statistic (see below) from observed
data was compared with those from 5000 randomizations
under the appropriate null model (see below), again using the
IV index (see above). Observed, expected and IV values were
calculated separately for each sub-quadrat. The significance
(P) of IV was calculated as the proportion of randomizations
giving an IV value equal to, or more extreme than, that
observed, adjusted to a two-tailed test (previous work has
shown departures from the null model in either direction,
and this is expected ecologically).
For constancy in species richness, the test statistic was the
variance in richness across the 100 10 10 cm sub-quadrats
of a quadrat. The null model comprised the number of
occurrences of each species in the quadrat being fixed at
that observed, but allocated to sub-quadrats at random,
Fig. 2. Heterogeneity among 10 10 cm sub-quadrats within a 1 1 m quadrat.
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907
separately for each species (Wilson et al. 1987). An overall
null model was used in which species occurrences were ran-
domized over all 100 sub-quadrats in a quadrat, and also a
patch model in which the null model for each target sub-
quadrat was based on a 3 3 sub-grid of sub-quadrats
around the target (Watkins & Wilson 1992).
For guild proportionality, the guilds used were forbs versus
all other species (there was no separation among the latter
because of the small number of species involved). The test
statistic was the variance in the proportion of species in the
sub-quadrat that were forbs, that is, the variance in (number
of forb species)/(total number of species) (Wilson & Watkins
1994). The guild proportions of the non-forb guild were not
analysed because they are the complement of the forb guild.
The null models, overall and patch, comprised holding the
species frequencies in each quadrat (with a site model) or
patch (with a patch model: Watkins & Wilson 1992) and the
richness of each sub-quadrat at those observed, but with
occurrences randomized within these constraints (Wilson
1989; using a swapping method of the type validated by
Mikls & Podani 2004).
RESULTS
Spatial community assembly
Spatial heterogeneity between sub-quadrats using
species presence/absence (Jaccard index; Fig. 2a) was
significantly higher at Coronet than at Cardrona
(F = 22.2, P < 0.001), but more notably the responses
to disturbance were quite different between the
two areas (area disturbance interaction F = 21.7,
P < 0.001). At Cardrona, the lowest presence/absence
heterogeneity was seen in the Ski-run, but at Coronet
heterogeneity was highest there (Fig. 2a).
When heterogeneity was weighted by species biom-
asses (abundance, index PD; Fig. 2b) the two areas
differed in the opposite way from presence/absence,
within-quadrat heterogeneity being higher at Card-
rona (F = 16.1, P < 0.001). Response to disturbance
again differed between the two areas, and in ways
different from that in presence/absence: at Cardrona
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908 F. B. J. C A S H ET AL.

the lowest heterogeneity was in the Adjacent, but at
Coronet it was highest there (interaction F = 3.54,
P = 0.043).
In summary, the response of heterogeneity to distur-
bance was inconsistent between the two areas, and
was also dependent on whether heterogeneity was
weighted by species biomasses.
Spatial autocorrelation within quadrats was signifi-
cant in the Adjacent and Undisturbed sites in both
areas (P < 0.01, by randomization test), but not in the
Ski-runs except at Coronet in terms of biomass (index
PD, P = 0.028).The strength of spatial autocorrelation
is indicated by examining the variation in dissimilari-
ties between quadrats in their species composition,
and calculating the percentage of it that is explained
by distance. This percentage differed significantly
between disturbance regimes, for presence/absence
(Jaccard; Fig. 3a, F = 5.16, P = 0.012) and for abun-
dance (PD; Fig. 3b, F = 3.83, P = 0.033). This differ-
ence between regimes was consistent between the two
areas sampled (area comparison P with Jaccard 0.84,
with PD 0.52; area disturbance interaction P with
Jaccard 0.20, with PD 0.66). In particular, spatial
autocorrelation was stronger (by Duncans test) in the
Undisturbed than in the Ski-run, with the Adjacent
generally being intermediate (Fig. 3). Thus, spatial
autocorrelation in the areas sampled was more pro-
nounced in the less-disturbed sites, as hypothesized.
Abundance-pattern assembly rules
Evenness at 10 10 cm was significantly higher in the
Ski-run sites than in the Adjacent or Undisturbed ones
(Fig. 4a, P < 0.022 for the disturbance effect).The two
areas showed similar, though not identical, effects (the
area disturbance interaction was non-significant,
P = 0.076), and this was not scale-dependent, for very
similar patterns were seen lumping adjacent quadrats
to 20 10 cm and 100 10 cm scales (results not
shown).
Of the RAD models, the Zipf-Mandelbrot gave the
best fit (i.e. it had the lowest sum of squares deviance)
in every case except Undisturbed at Cardrona, for
which it was second-best to General Lognormal
(Table 1).The Zipf-Mandelbrot fit was especially close
for the two Ski-run sites. Broken Stick did not give the
best fit for any individual quadrat, or for any site (the

Fig. 3. Spatial autocorrelation (SAC): the percentage of variation in species composition explained by distance: (a) for
presence/absence, measured by Jaccard coefficient (b) for abundance (biomass), measured by proportional dissimilarity
coefficient.

Fig. 4. (a) Evenness within among 10 10 cm sub-quadrats. (b) Rank consistency index compared to null model values with
an IV index. High values indicate that the biomass rank order of species is high. 0, observed is equal to null model, +, observed
is greater.The absolute residuals were correlated with the fitted (expected) values (r = +0.39), but the assumptions of anova were
otherwise met.

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Table 1. Best fits of species biomass to models of relative abundance distribution

Best fit Second-best fit

Area Site Model SS deviance Model SS deviance

Cardrona Ski-run Zipf-Mandelbrot 0.087 Geometric 0.327

Adjacent Zipf-Mandelbrot 0.229 General lognormal 0.509
Undisturbed General lognormal 0.268 Zipf- Mandelbrot 0.424
Coronet Ski-run Zipf-Mandelbrot 0.016 General lognormal 0.635
Adjacent Zipf-Mandelbrot 0.072 General lognormal 0.337
Undisturbed Zipf-Mandelbrot 0.093 Geometric 0.771

For each separate sub-quadrat, the deviances are scaled so that the best-fitting model has a sum of squares deviance (SS
deviance) of 0.0.

Table 2. Constancy in richness: variance in richness compared to a null model with an IV index

Area Cardrona Coronet

Overall model Patch model Overall model Patch model

Disturbance IV P No. sig IV P No. sig IV P No. sig IV P No. sig

Ski-run -1.8 ns 0 +1.2 ns 0 +0.1 ns 1/6 +2.3 ns 0

Adjacent +11.1 ** 3/6 +6.1 * 1/6 -7.6 * 2/6 -2.5 ns 0
Undisturbed -3.0 ns 1/6 +0.6 ns 2/6 +6.2 * 2/6 +3.6 ns 0

*<0.05; ***<0.001. 0, observed is equal to null model; +ve, observed is greater; -ve, observed is less (an assembly rule); ns, not
significant; No. sig, the number of the six sites showing significant effects on their own.

Broken Stick model implies very high evenness). For
most sites, the second-best fit was Geometric or
General Lognormal; in fact, overall the fits of these two
models were very close (Geometric average = 0.571,
General Lognormal = 0.584), and there was no ten-
dency for a particular model (Broken Stick, Geomet-
ric, General Lognormal and Zipf-Mandelbrot) to fit
better in one of the three disturbance regimes.
As expected, some rank consistency was always
present, as indicated by IV values being positive (there
should be some consistency among the 10 sub-
quadrats of a quadrat in which species have high
biomass and which have low biomass). It was signifi-
cantly greater than the null model in 34 out of the
36 quadrats (3 disturbance regimes 2 areas 6
replicates). There was no overall difference between
disturbance states in this consistency in rank
(P = 0.66), but the response to disturbance at the two
areas was quite different: at Cardrona the Ski-run had
the lowest rank consistency and at Coronet it had
the highest (Fig. 4b, area disturbance interaction
P = 0.019).
ness from that expected under the null model and with
some departures in each direction (Table 2). In the
Coronet Adjacent sites using an overall model, rich-
ness was significantly more constant than expected
(i.e. IV < 0), as would be expected from niche
limitation. However, once environmental heterogene-
ity and spatial autocorrelation effects were excluded by
using a patch model, the effect, though still present
(IV = -2.5), was no longer significant. The only effect
remaining significant under a patch model was high
variance in richness in the Cardrona Adjacent sites,
suggesting environmental variation rather than an
assembly rule (IV = +6.1).
Guild proportionality was strong and highly signifi-
cant (i.e. lower variation in the proportional represen-
tation of the forb guild than expected under a null
model, IV < 0.0) in the Ski-run sites in both areas
(Table 3). However, these effects decreased and
became non-significant using a patch model.
DISCUSSION

Niche-based assembly rules
Constancy in species richness was not generally appar-
ent, with only sporadic departure of variance in rich-
2012 The Authors
Austral Ecology 2012 Ecological Society of Australia
There have been many studies of communities under
different degrees of, or different times since,
disturbance. Few generalizations have been possible,
but studies of spatial and assembly patterns offer that
doi:10.1111/j.1442-9993.2011.02349.x
910 F. B. J. C A S H ET AL.
Table 3. Guild proportionality: variance in proportion of species in the forb guild compared to a null model with an IV index
Area Cardrona
Overall model
Disturbance IV P No. sig IV
Ski-run -7.3 0.0028 1/6 -1.1
Adjacent +5.2 0.0084 4/6 +1.4
Undisturbed -4.2 ns 0 +0.8
The two quadrats with significance showed it in opposite directions. 0, observed is equal to null model; +ve, observed is
greater; -ve, observed is less (an assembly rule); ns, not significant; No. sig, the number of the six sites showing significant effects
on their own.
opportunity. In order to avoid drawing conclusions
from only one area, we chose two different but com-
parable ski fields. However, this still represents only
minimal replication at the area level. We can therefore
draw conclusions only about the two particular areas.
Thus, although differences between the sites are good
evidence of inconsistent response to disturbance,
similar responses cannot be taken as evidence that
there would be such a response in other areas. Simi-
larly, we can draw conclusions only about the spatial
grain sampled, as in the great majority of ecological
studies, although quadrats sizes were chosen to be
appropriate for the communities.
Trends in heterogeneity
Greig-Smith (1952) and Gitay and Wilson (1995) sug-
gested that there would be three phases in succession
after a disturbance: (i) Pioneer, with moderate hetero-
geneity resulting from random initial colonization; (ii)
Building, with reduced heterogeneity as species
spread; and (iii) Mature, with strong heterogeneity
arising from micro-habitat sorting and the operation of
assembly rules. Gitay and Wilson (1995) found
support for this model in the recovery of New Zealand
tussock grasslands from fire.The observation of Glenn
et al. (1992) also matches the model: in prairie sites
that had not been experimentally disturbed by grazing
and/or burning (i.e. the Mature phase) there was
higher heterogeneity. Abundance-based heterogeneity
at Cardrona (Fig. 2b) exactly matched the theory,
higher in the Ski-run (Pioneer phase) than in the Adja-
cent vegetation (Building phase), then higher again in
the Undisturbed (Mature phase). However, the oppo-
site pattern was seen at Coronet. Apart from the higher
heterogeneity in the Ski-run at Coronet, presence/
absence heterogeneity did not fit the theory either
(Fig. 2a).The differences between the two sites cannot
be explained in terms of known disturbance history, as
it is clear that both were originally denuded to bare soil
doi:10.1111/j.1442-9993.2011.02349.x
Coronet
Patch model Overall model Patch model
P No. sig IV P No. sig IV P No. sig
ns 0 -8.6 <0.001 2/6 -0.8 ns 0
ns 0 -1.1 ns 2/6 +0.2 ns 0
ns 0 -1.8 ns 1/6 +1.9 ns 0
and then colonized from sown and volunteer seed.
Some reports in the literature conflict with the theory
too. Seabloom et al. (2005) studied experimental
grasslands in California over the 4 years after sowing
or burning.This timespan surely represents the span of
the Pioneer to the Building phases, when heterogene-
ity should decrease by the theory, but instead it
increased. Ruprecht et al. (2007) found in a chronose-
quence of old fields in Romania that there were no
trends in the proportion of statistically significant
species associations, contrary to the theory. Lepori and
Malmqvist (2009), investigating freshwater inverte-
brates, found the highest heterogeneity under the least
disturbed conditions and the lowest under intermedi-
ate disturbance, in conformance with theory. It seems
that changes in community heterogeneity after distur-
bance are not simple: the pattern varies between areas,
as it did here, and different measures of species com-
position need to be examined to check that they are
consistent. Much of the evidence is from space-for-
time substitution studies, and more studies of tempo-
ral changes within individual sites are needed.
Biomass heterogeneity (Fig. 2b) comprises two
aspects: which species are present (Fig. 2a) and
whether they are present in consistent amounts (Fig.
4b).The latter aspect is rank consistency, or rather the
inverse of it because the greater the rank consistency
the lower the abundance heterogeneity. The results
for rank consistency (Fig. 4b) do indeed closely mirror
those for abundance heterogeneity (Fig. 2b). The
unexpected Coronet result, of high rank consistency in
the ski-run, best matches the observation of Collins
et al. (2008), of higher rank consistency with higher
burning disturbance in prairie. For the ski-run, this
may have been because the species present were pre-
dominantly exotic, with efficient dispersal, but it is not
clear whether this applies to the burnt prairie.
Spatial differences are not just a question of the
amount of heterogeneity, but also of the consistency of
spatial trends in that heterogeneity, that is, whether the
dissimilarity in community composition increases
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 A S S E M B LY R U L E S I N D I S T U R B E D C O M M U N I T I E S
steadily with distance apart. This is spatial auto-
correlation. The study of spatial autocorrelation
at the community level is relatively new (Kleb &
Wilson 1999; Nekola & White 1999; Mistral et al.
2000) and little is known of its change through succes-
sion. We would expect the strength of spatial autocor-
relation to increase through succession as the
community becomes organized, and it generally did
(Fig. 3). In the only comparable study, Lu et al. (2009)
reported for four Chinese wetlands stronger spatial
autocorrelation in those with lower disturbance, indi-
cated by a lower agricultural ditch density in the sur-
rounding lands. This parallels our result, and may be a
general rule in succession, but observations over time
are needed.
In summary, the prediction of Fowler (1990) that
disturbance would increase the role of disorderliness/
random-events receives only limited support in terms
of pure heterogeneity, but it is strongly supported by
spatial autocorrelation: the strength of the fitted
dissimilarity/distance relation increased as the amount
of disturbance decreased. Although the amount of het-
erogeneity cannot be predicted, its spatial distribution
can.
Trends in abundance patterns
The evenness of a community is a very obvious result
of species interactions and assembly: the simplest
abundance-based assembly rule. It represents species
dominance, that is, heterogeneity among the species.
There is no theory on its response to disturbance,
but many empirical studies. Mackey and Currie
(2001) found that in 50% of published studies there
was no significant trend, but of the remainder the
majority indicated that disturbance reduces evenness.
However, almost all such studies, those reviewed by
Mackey and Currie and those published since, are
flawed in that: (i) the measure of plant abundance
has usually been subjective, or even not stated; or (ii)
an evenness index such as J has been used that
reflects species richness as well as evenness (Smith &
Wilson 1996) and usually species richness did change
with disturbance in the studies Mackey and Currie
reviewed. In a study of two successional sequences
using biomass and the richness-independent index
E, Wilson et al. (1996) showed evenness to increase
through succession, but the present work with the
same methods found the opposite pattern: evenness
was lower in the less-disturbed communities
(Fig. 4a). No generalization is yet possible. Evenness
will depend on the relative competitive ability of the
species and the opportunity for niche complementa-
rity, so successional trends in evenness as in the
Wilson et al. (1996) study, or differences between
sites with difference disturbance histories as here,
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Austral Ecology 2012 Ecological Society of Australia
911
may be individualistic to each site with its unique
assemblage of species.
A more subtle abundance-based assembly rule is
the RAD. Community models have predicted that a
lognormal RAD will be found at equilibrium but not
in disturbed communities (Stenseth 1979; Ugland &
Gray 1982). Similarly, in the theoretical study of
Wissel and Maier (1992) only those models that
included competition, such as would predominate
later in succession, gave lognormal distributions.
Some animal ecologists have claimed to find confir-
matory evidence that a lognormal distribution is
typical of, and restricted to, equilibrium communities
(e.g. Hamer et al. 1997), but others have failed to see
a general pattern (Nummelin 1998). Evidence for
plant communities is very sparse. Wilson et al. (1996)
could find no consistent pattern of change in RAD
model fit through succession. In the present results,
the General Lognormal did give the best fit to the
Undisturbed vegetation at Cardrona, conforming to
the theory, but for the Undisturbed at Coronet it
was inferior to both the Zipf-Mandelbrot and the
Geometric. Insight into the cause of abundance dis-
tributions can also be found via rank consistency
since the frequent comments that disturbance will
cause founder events to predominate and prevent
communities from self-organizing (Fowler 1990;
Drake et al. 1999) imply that, in the absence of dis-
turbance, evolutionary/competition control will be
present, giving higher rank consistency (Watkins &
Wilson 1992). This prediction was borne out at
Cardrona, but the opposite was true at Coronet
(Fig. 4b). Again, any generalizations would be
premature.
The conclusion on abundance-based assembly rules
must be that there are no trends consistent between
sites, or that conform to theoretical predictions.
Niche-based assembly rules
Assembly rules such as constancy in species richness
and guild proportionality suggest that different
patches of a community will converge in structure,
even if they differ in species composition. The rules
will be only trends because of the noise of incomplete
competitive exclusion, overlap between niches, empty
niches, etc., but they may still be detectable when
compared to a null model in which no niche constraint
is present (Wilson et al. 1987; Wilson 1989). We
started with the widely held concept that this would be
true only for equilibrium communities (Bartha et al.
1995; Belyea & Lancaster 1999; Cornell 1999; Drake
et al. 1999). Part of the reasoning has been that after a
disturbance competitive intensity is reduced (Bazzaz
1987; Wilson & Tilman 1993). Violle et al. (2010)
doi:10.1111/j.1442-9993.2011.02349.x
912 F. B. J. C A S H ET AL.
have questioned this, but the logic remains that time is
required for species to disperse, exclusion to occur,
and abundances to adjust.
Only a little evidence has been produced to
support this. Wilson et al. (1987) and Wilson and
Gitay (1995) demonstrated that the least disturbed of
four dune slacks was the one showing evidence of
niche limitation and guild proportionality. In the
present work the only significant indication of con-
stant richness (negative IV) was in the Adjacent sites
at Coronet (Table 2), while guild proportionality was
evident in both Ski-run sites (Table 3): the opposite
position in the disturbedundisturbed spectrum from
that expected by the logic of Belyea and Lancaster
(1999) and others. One possible explanation for the
guild proportionality results is that mutualisms are
important early in the ski-runs, which are in an early
successional stage after disturbance, as after distur-
bance in other alpine areas (Choler et al. 2001),
giving the relative constancy in guild proportions
seen in the ski-runs (Tables 2,3). In all three cases
the effects became small and non-significant using a
patch model, so the cause may be environmental
micro-heterogeneity (Wilson 1999). Most other
studies of constancy in richness and guild propor-
tionality have not used a patch model, and their
results may likewise reflect habitat sorting. However,
we must bear in mind that our effects might be real,
but not demonstrable with a patch model, as basing
the null for a target quadrat on only the nine sur-
rounding quadrats gives a conservative test.
Conclusions
The hypothesis with which we started was stated
rather explicitly by Fowler (1990), that disturbance
produces disorderliness in plant communities: it
weakens the effectiveness of both the processes that
promote species coexistence and also those that
cause competitive exclusion, so that random events
are more important. This affects the frequency with
Fig. 5. Spatial autocorrelation: the nugget (intercept, interpreted as a measure of intrinsic randomness: (a) for presence/
absence, measured by Jaccard coefficient (b) for abundance (biomass), measured by proportional dissimilarity coefficient.
doi:10.1111/j.1442-9993.2011.02349.x
which the outcome of competition (winner vs. loser)
can be identified. Our results come close to disprov-
ing this as a general phenomenon. Higher evenness
was seen in disturbed site at Coronet, but this is con-
trary to trends observed elsewhere (Wilson et al.
1996), and there was no other abundance-based
assembly rule. The only niche-based assembly rules
might have been caused by environmental differences
between 10 10 cm patches since they disappear
with a patch model. Spatially, the lack of consistency
between whether heterogeneity was higher or lower
with disturbance reflects mixed reports in the litera-
ture, and indeed mixed prediction from theory.
The one consistent pattern is stronger spatial auto-
correlation in undisturbed sites, supporting the
concept of the Mature phase in the Greig-Smith/
Gitay and Wilson model of community assembly:
decreased randomness but increased patchiness,
reflecting not only micro-habitat sorting but also
interspecific interactions of the type that would cause
assembly rules.
In the Cardrona area, more consistent support can
be seen for the arguments of Bartha et al. (1995),
Belyea and Lancaster (1999) and Drake et al. (1999).
There is low initial (Pioneer phase) heterogeneity in
species presence since any species can arrive, but
high biomass patchiness due to founder dominance
(Fig. 2). Then, after a homogeneous intermediate
Building phase, the final Mature phase has high het-
erogeneity in both presence and biomass due to
habitat and competitive sorting (Fig. 2). This is
accompanied by steadily increasing spatial autocorre-
lation, as expected (Fig. 3), although unexpectedly
the nuggets were lower in the ski-runs (Fig. 5). There
is little theory to guide us on the development of
evenness, but the evidence so far suggests that it will
increase through succession, and at least the opposite
trend is not seen at Cardrona (Fig. 4a). Several
models suggest that equilibrium communities should
have a general lognormal RAD, and this is seen at
Cardrona (Table 1). The expected shift from founder
control to dominance control, giving an increase in
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 A S S E M B LY R U L E S I N D I S T U R B E D C O M M U N I T I E S
rank consistency from disturbed to undisturbed, is
seen there (Fig. 4b). The analyses of constancy in
richness (Table 2) and guild proportionality (Table 3)
are dominated by the effects of patchiness. Neither
show unequivocally the assembly rules predicted, but
this is unsurprising because such rules are difficult to
demonstrate (Wilson et al. 1987; Wilson 1989,
1999). The evidence for Cardrona points to the pro-
cesses developing that lead to assembly rules, even if
the end results are obscured.
However, the absence of many of these patterns at
Coronet reminds us that nature does not always neatly
follow our theories. More empirical studies are needed
in order to establish general trends.
ACKNOWLEDGEMENTS
We thank the land holders for access, especially Mr
and Mrs R.E. Anderson for accommodation and facili-
ties, Prof. A.F. Mark for advice, Vickey Tomlinson for
help in sorting plant material and John Steel for com-
ments on a draft.
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